ISSN 0006 8241 = Bothalia

Bothalia

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Vol. 23,1

May/Mei 1993

PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE, PRETORIA

PUBLIKASIES VAN DIE NASIONALE BOTANIESE INSTITUUT, PRETORIA

Obtainable from the National Botanical Institute, Private Bag X101,

Pretoria 0001, Republic of South Africa. A catalogue of all available

publications will be issued on request.

BOTH A LI A

Bothalia is named in honour of General Louis Botha, first Premier

and Minister of Agriculture of the Union of South Africa. This

house journal of the National Botanical Institute, Pretoria, is devoted to

the furtherance of botanical science. The main fields covered are

taxonomy, ecology, anatomy and cytology. Two parts of the journal

and an index to contents, authors and subjects are published

annually.

Verkrygbaar van die Nasionale Botaniese Instituut, Privaatsak X101,

Pretoria 0001, Republiek van Suid-Afrika. ’n Katalogus van alle beskik-

bare publikasies kan aangevra word.

Bothalia is vemoem ter ere van Generaal Louis Botha, eerste Eerste

Minister en Minister van Landbou van die Unie van Suid-Afrika. Hierdie

lyfblad van die Nasionale Botaniese Instituut, Pretoria, is gewy aan die

bevordering van die wetenskap van plantkunde. Die hoofgebiede wat gedek

word, is taksonomie, ekologie, anatomie en sitologie. Twee dele van die

tydskrif en ’n indeks van die inhoud, outeurs en onderwerpe verskyn

jaarliks.

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

MEMOIRS VAN DIE BOTANIESE OPNAME VAN SUID-AFRIKA

The memoirs are individual treatises usually of an ecological nature, ’n Reeks van losstaande omvattende verhandelings oor vemaamlik

but sometimes dealing with taxonomy or economic botany. ekologiese, maar soms ook taksonomiese of plantekonomiese onderwerpe.

THE FLOWERING PLANTS OF AFRICA / DIE BLOMPLANTE VAN AFRIKA

This serial presents colour plates of African plants with accompanying

text. The plates are prepared mainly by the artists at the National Botanical

Institute. Many well known botanical artists have contributed to the series,

such as Cythna Letty (over 700 plates), Kathleen Lansdell, Stella Gower,

Betty Connell, Peter Bally and Fay Anderson. The Editor is pleased to

receive living plants of general interest or of economic value for

illustration.

From Vol. 50, one part of twenty plates is published annually. A volume

consists of two parts. The publication is available in English and Afrikaans.

Hierdie reeks bied kleurplate van Afrikaanse plante met bygaande teks.

Die skilderye word meestal deur die kunstenaars van die Nasionale

Botaniese Instituut voorberei. Talle bekende botaniese kunstenaars het

tot die reeks bygedra, soos Cythna Letty (meer as 700 plate), Kathleen

Lansdell, Stella Gower, Betty Connell, Peter Bally en Fay Anderson.

Die Redakteur verwelkom lewende plante van algemene belang of

ekonomiese waarde vir afbeelding.

Vanaf Vol. 50 word een deel, bestaande uit twintig plate, jaarliks

gepubliseer. ’n Volume bestaan uit twee dele. Die publikasie is beskikbaar

in Afrikaans en Engels.

FLORA OF SOUTHERN AFRICA / FLORA VAN SUIDELIKE AFRIKA

A taxonomic treatise on the flora of the Republic of South Africa,

Ciskei, Transkei, Lesotho, Swaziland, Bophuthatswana, Namibia,

Botswana and Venda. The FSA contains descriptions of families, genera,

species, infraspecific taxa, keys to genera and species, synonymy, literature

and limited specimen citations, as well as taxonomic and ecological

notes.

’n Taksonomiese verhandeling oor die flora van die Republiek van Suid-

Afrika, Ciskei, Transkei, Lesotho, Swaziland, Bophuthatswana, Namibie,

Botswana en Venda. Die FSA bevat beskrywings van families, genusse,

spesies, infraspesifieke taksons, sleutels tot genusse enspesies, sinonimie,

literatuur, verwysings na enkele eksemplare, asook beknopte taksonomiese

en ekologiese aantekeninge.

PALAEOFLORA OF SOUTHERN AFRICA / PALAEOFLORA VAN SUIDELIKE AFRIKA

A palaeoflora on a pattern comparable to that of the Flora of

southern Africa. Much of the information is presented in the form

of tables and photographic plates depicting fossil populations. Now

available:

’n Palaeoflora met ’n uitleg vergelykbaar met die van die Flora van suide-

like Afrika. Baie van die inligting word aangebied in die vorm van tabelle

en fotografiese plate waarop fossiele populasies afgebeeld word. Reeds

beskikbaar:

Molteno Formation (Triassic) Vol. 1. Introduction. Dicroidium , by/deur J.M. & H.M. Anderson.

Molteno Formation (Triassic) Vol. 2. Gymnosperms (excluding Dicroidium), by/deur J.M. & H.M. Anderson.

Prodromus of South African Megafloras. Devonian to Lower Cretaceous, by/deur J.M. & H.M. Anderson.

Obtainable from/Beskikbaar van: A. A. Balkema Marketing, Box/POsbus 317, Claremont 7735, RSA.

BOTHALIA

N TYDSKRIF VIR PLANT KUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Volume 23,1

Scientific Editor/Wetenskaplike Redakteur: O.A. Leistner

Technical Editor/Tegniese Redakteur B.A. Momberg

NATIONAL

INSTITUTE

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag XI01, Pretoria 0001

ISSN 0006 8241

. -G5- i 1

Privaateak X101 PRETORIA 00'v

national botanical.

INSTITUTE

J

1993

Editorial Board/Redaksieraad

D.E Cutler

B.J. Huntley

P.H. Raven

J.P. Rourke

M.J. Werger

Royal Botanic Gardens, Kew, UK

National Botanical Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

Compton Herbarium, NBI, Cape Town, RSA

University of Utrecht, Utrecht, Netherlands

Editorial Committee

Redaksiekomitee

O.A. Leistner

B.A. Momberg

M.C. Rutherford

CONTENTS — INHOUI)

Volume 23,1

1 . Studies in the Ericoideae (Ericaceae). XU. The placing of the genus Blaeria into synonymy under Erica ;

nomenclatural and taxonomic changes for the southern African region. E.G.H. OLIVER 1

2 . Studies in the Ericoideae (Ericaceae). Xm.. Three new species of Erica from the southwestern Cape.

E.G.H. OLIVER 9

3. The hepatics, Symphyogyna podophylla and Pallavicinia lyellii (Pallaviciniaceae) in southern Africa.

S.M. PEROLD 15

4. A biosystematic study of Pentameris (Arundineae, Poaceae). N.P. BARKER 25

5 . Studies in the Marchantiales (Hepaticae) from southern Africa. 1. The genus Dumortiera and D. hirsu-

ta; the genus Lunularia and L. cruciata. S.M. PEROLD 49

6. Panicum simulans (Paniceae, Poaceae), a new species from southern Africa and its leaf anatomy. L.

SMOOK and R.P. ELLIS 59

7. Notes on African plants:

Allisoniaceae. The hepatic, Calycularia crispula (Metzgeriales) reported from Malawi and

Zambia. S.M. PEROLD 79

Asteraceae. An evaluation of Hutchinson’s ‘beetle-daisy’ hypothesis. J.J. MIDGLEY 70

Fabaceae. Vigna kokii, a new species from southern Africa. BJ. PIENAAR 68

Fabaceae. Notes on the genus Argyrolobium (Crotalarieae) including a new species from southern

Africa. T.J. EDWARDS 77

Oxalidaceae. A new species of Oxalis from the western Cape. E.G.H. OLIVER 72

Pteridophyta— Adiantaceae. A new cytotype for Acrostichum aureum. J.P. ROUX 75

Rosaceae. Observations on Cliffortia micrantha. A.C. FELLINGHAM 65

Rosaceae. Cliffortia fasciculata, a superfluous name for C. amplexistipula. A.C. FELLINGHAM 67

8. First report on the presence of Enterobryus species (Trichomycetes: Eccrinales) in South Africa and

the description of three new species. G.J.M.A. GORTER 85

9. Mycorrhizal status of plants growing in the Cape Floristic Region, South Africa. N. ALLSOPP and

W.D. STOCK 91

10. Pollen morphology of Curroria, Mondia, Socotranthus and Stomatostemma (Periplocaceae) . R.L.

VERHOEVEN and H.J.T. VENTER 105

11. Dynamics of the forest vegetation of the Umtiza Nature Reserve, East London. J.J. MIDGLEY and

P.N. GOBETZ Ill

12 .The vegetation of the northeastern Orange Free State, South Africa: physical environment and plant

communities of the Ea land type. H.C. ECKHARDT, N. VAN ROOYEN and G.J. BREDEN-

KAMP 117

13 . The vegetation of the southern Langeberg, Cape Province. 1. The plant communities of the Boosmans-

bos Wilderness Area. D.J. MCDONALD 129

14. The vegetation of the southern Langeberg, Cape Province. 2. The plant communities of the Marloth

Nature Reserve. D.J. MCDONALD 153

Digitized by the Internet Archive

in 2016

https://archive.org/details/bothaliavolume2323unse

Bothalia 23,1: 1-7 (1993)

Studies in the Ericoideae (Ericaceae). XII. The placing of the genus

Blaeria into synonymy under Erica ; nomenclatural and taxonomic

changes for the southern African region

E.G.H. OLIVER*

Keywords: Blaeria, Erica, nomenclature, southern Africa, taxonomy

ABSTRACT

The reduction of the genus Blaeria to synonymy under Erica requires the publication of six new combinations and three

new names for the nine taxa occurring in southern Africa. Four species are reduced to synonymy. Diagnostic features and

distribution maps are provided.

UITTREKSEL

Die plasing van die genus Blaeria in sinonimie onder Erica vereis die publisering van ses nuwe kombinasies en drie nuwe

name vir die nege taksons wat in suidelike Afrika voorkom. Vier spesies word in sinonimie geplaas. Diagnostiese kenmerke

en verspreidingskaarte word voorsien.

INTRODUCTION

In a forthcoming paper (Oliver in press A) the generic

relationship between Blaeria L. and Erica L. will be

discussed in detail. The former genus has been maintained

separate from Erica since 1753 on the single character

difference of four as opposed to eight stamens.

Problems arose with several species in both genera

having a variable number of stamens or even numbers not

included within the circumscription of their genera.

Depending on the stamen complement found in the flowers

examined, one could identify some material as either

belonging to Blaeria or to Erica. Problem species in this

respect are Erica filiformis Salisb., E. esterhuyseniae

Compton and E. pleiotricha S. Moore in which 8- and

4-stamened flowers exist. In several other species the

stamen number varies from 5 to 8.

With the sinking of the genus Philippia into Erica

(Oliver 1987a, 1988, 1989) several additional species from

tropical Africa having variable numbers of stamens were

introduced into the problem. These are: E. rryassana (Aim

& Fries) E.G.H. Oliver with only 4 stamens, very occa-

sionally 5; E. hexandra (S. Moore) E.G.H. Oliver with

6, but 7 or 8 in some or all flowers; E. mannii (Hook,

f.) Beentje has 6 stamens but with 5, 7 or 8 in some

flowers. Several Madagascan species, not yet transferred

to Erica, have also compounded the problem; Philippia

humbertii H. Perrier with 3 or 4 stamens, sometimes 6

and P gracilis (Benth.) H. Perrier with 4-8 stamens.

The genus Blaeria contains approximately 16 species,

all occurring in Africa. In the southern African region the

nine species are confined to the Cape. They can be placed

in several groups of related species, with each group

showing more affinities to different sections within Erica

* Stellenbosch Herbarium, National Botanical Institute, P.O. Box 471,

Stellenbosch 7599.

MS. received: 1992-06-09.

than to the other groups within Blaeria, including those

from tropical Africa.

It is postulated (Oliver in press A) that Blaeria is an

unnatural genus with clear indications of being poly-

phyletic. On the grounds of polyphylesis and the complete

transition between the two genera in the only differentiat-

ing character in certain species, it was decided to reduce

the genus Blaeria to synonymy under Erica. This decision

is formalised in this paper for the species occurring in the

southern African region. At this stage it is not possible

to deal with the nomenclatural changes necessary for the

tropical African species because of the instability of their

taxonomy and of several nomenclatural synonyms within

Erica.

1. Erica barbigeroides E.G.H. Oliver, nom. nov.

Blaeria revoluta Bartl.: 650 (1832), non Erica revoluta (H. Bol.) L.E.

Davidson (1985); Klotzsch: 663 (1833); Klotzsch: 222 (1838); Benth.:

698 (1839); N.E. Br.: 326 (1905). Blairia revoluta (Bartl.) Dietr.: 444

(1839). Type: Kleinriviersbergen, Ecklon s.n. (Bt, holo.; S!). Lectotype

chosen here: Ecklon s.n. [det. Bartl.] (S).

Blaeria barbigera sensu Aim & Fries: 235 (1924); Bond & Goldblatt:

239 (1984); Oliver: 145 (1987b).

This very distinct species was referred to as Blaeria

barbigera based only on the assumption that the material

fitted the description of Salisbury’s Erica barbigera (1802),

there being no extant type specimen. For further details

see below under Insufficiently known species.

The new name is an adaptation of the name by which

this well-known species of the coastal regions of the

Caledon District has been known for many years.

The species is confined to sandy level areas that are often

wet in winter in the coastal region of the southwestern Cape

from Rooi Els in the west to Sondagskloof (Sandies Glen)

near Napier in the east (Figure 1). In this region it may

be found from just above the spray zone near the sea up

to an altitude of 760 m on the nearby mountains.

2

18" 19° 20° 21°

20 0 20 40 60 80 100 km

FIGURE 1. — The known distribution of Erica barbigeroides.

E. barbigeroides forms sparsely branched, virgate

shrublets up to 400 mm tall, is covered with numerous

long hairs and has up to 24 flowers grouped together

in closely packed heads at ends of branches. Heads

often partially pendent. Flowers somewhat sticky due to

secretion of viscid matter from sessile glands on margins

of sepals.

It is allied to two other species mentioned here, E.

ericoides and E. russakiana, both of which occur in the

same area near Hermanus, but never grow together. Both

of the latter species form many-branched shrublets and are

smaller in all parts of the inflorescence and flower.

Vouchers: Ecklon & Zeyher 260 [loc. 58.8] (BOL, G, MO, S, SAM,

W); Ecklon <6 Zeyher s.n. [loc. 58.8] (K, LD, P, UPS, W, Z); Oliver

& Palser 83 (E, MEL, NY, PRE, STE); Schlechter 95U (BM, BOL,

G, K, MO, PRE, STE, W, Z). 70 other collections examined.

2. Erica equisetifolia Salisb. in Transactions of the

Linnean Society 6: 342 (1802). Type: locality & collector

unknown, Herb. Salisb. (K, holo.!).

Blaeria equisetifolia (Salisb.) G. Don: 805 (1834); Aim & Fries: 239

(1924); Salter: 658 (1950); Bond & Goldblatt: 239 (1984); Oliver: 145

(1987).

B. purpurea L. f.: 122 (1782) nom. illegit. non Berg. (1767) [= Simo-

cheilus purpureus (Berg.) Druce]; Thunb.: 8 (1802); Klotzsch: 221 (1838);

N.E. Br. : 323 (1905); Aim & Fries: 239 (1924) pro parte. Type: Thunberg

s.n. (UPS, holo.!).

B. dumosa Wendl.: t .38 (1808); Roem. & Schult.: 170 (1818); G. Don:

805 (1834); N.E. Br.: 323 (1905), synon. nov. Type: Wendl.: t.38.

B. dumosa var. breviflora N.E. Br.: 323 (1905), synon. nov. Syntypes:

Caledon Div., mountains near Genadendal, Bolus 5419 (BOL!, K!, PRE!)

and in Herb. Norm 613 (K!); ibid. Guthrie 3140 (?); without locality,

Drege s.n. (K!) .

B. campanulata Benth,: 698 (1839); N.E.'Br. : 324 (1905), synon. nov.

Syntypes: Cape, Drige s.n. (?); Burchell 7693 (K!); Burchell 7773 (K!,

P!, W!).

B. flava H. Bol.: 239 (1894); N.E. Br.: 322 (1905), synon. nov. Syn-

types: Zwartberg near Caledon, Jan. 1885, 800 m, Bolus 5147 (BOL!,

Bothalia 23,1 (1993)

K!, PRE!, STE!) & sub Herb. Norm. 611 (BM!, BOL!, G!, NH!, P!,

PRE!, SAM!, UPS!, W!).

Erica parvula Guth. & Bol.: 171 (1904), synon. nov. Type: Stellenbosch

Div. ; on a rock near the mouth of the Steenbrass River, 20—30 ft. above

the sea, Guthrie 3710 (BOL, holo.!).

Blaeria oppositifolia L. Guthrie: 21 (1928), synon. nov. Type: Hotten-

tots Holland Mountains, Jan. 1924, Stokoe in BOL 17674 (BOL, holo.!,

K!).

Common and widespread in mountains of southwestern

Cape from Cape Peninsula to Bain’s Kloof in north to

western part of Bredasdorp District in south and occurs

from sea level to summits of mountains at 1 600 m (Figure

2). Highly variable species in habit, flower size and shape,

anther form and in having some collections with more than

usual number of four stamens.

Basic type forms rounded low shrublet with deep pink,

narrowly tubular flowers with four dark brown, exserted

anthers. In some cases, mainly at lower altitudes, plants

can be erect and reach 500 mm in height when growing

in old fynbos. Flowers may be short with corolla 2.5 mm

long and open campanulate to large with corolla 4 mm

long and tubular.

Several species formerly recognised as distinct have been

reduced to synonymy under E. equisetifolia on the grounds

of overlap in characters caused by the variation found in

the numerous collections of this complex. E. equisetifolia

and Blaeria dumosa were distinguished on the single

character of anther shape, the former having straight-sided

anthers whereas the latter had anthers that were obtriangu-

lar in outline with a constriction above the decurrent

appendages. The straight-sided collections come mostly

from the Cape Peninsula and the other material from the

inland mountains. There are collections from the main-

land which fit equally well in either of the two species.

Salisbury’s type is a small branchlet with ‘Equisetifolia

MS’ written in his own hand. It possesses six flowers with

only a single stamen remaining. From this stamen it is

Bothalia 23,1 (1993)

3

seen that the anther is the straight-sided form, most

probably from the Cape Peninsula, and that there is only

one small awn on one side.

Blaeria flava was described by Bolus on the grounds

of the yellow colour of the corolla. This would appear to

be the only character showing any discontinuity in this

complex which has only pink or white flowers. The species

is known only from the original collection from the

Swartberg at Caledon where no pink- or white-flowered

collections have been made. A thorough investigation of

the Swartberg may produce more evidence for assigning

this material some taxonomic status.

The problem of the similarity between E. equisetifolia,

Blaeria camparullata and E. parvula is mentioned (Oliver

in press A) as a case for reducing Blaeria to synonymy

under Erica.

B. campanulata was based on material collected from

rock ledges on the cool, southeast side of the top of the

mountains of Baviaanskloof above Genadendal from where

it is known through only a few collections. Plants are small

and very compact, the white flowers have exserted dark

stamens which range from 5—8 per flower and mostly

3-locular ovaries ( Oliver 8978 & 9081). A recent collec-

tion ( Oliver 8813 ) with only 8-stamened flowers has been

made on the lowest dry northern slopes near McGregor.

This material, in turn, was found to be morphologically

inseparable from E. parvula which is confined to the

Elgin/Betty’s Bay area near the coast. This latter species

also forms compact woody shrublets with white flowers.

It grows mostly on rocks in streambeds. Apart from the

more variable number of stamens and ovary locules the

material of these two species is indistinguishable. There

could, however, be some purpose in recognizing the high

altitude form from Baviaanskloof as a subspecies. This

will need some further investigations and assessment.

Several new species from the Kogelberg and Hermanus

areas that are closely allied to this species are being

described separately (Oliver 1993). One of these is re-

corded as producing a putative natural hybrid with E.

equisetifolia .

An anomalous form with mostly opposite leaves was col-

lected by Stokoe somewhere in the Hottentots-Holland

Mountains. Sometimes one can find an occasional 2-nate

arrangement of leaves on a branchlet in E. equisetifolia,

and in other species, such as the E. tenuifolia L.IE. lutea

Berg, complex, whole branchlets or even branches can

have 2-nate leaves. This condition alone is not regarded

as sufficiently distinct to warrant taxonomic recognition,

especially as it is known only from a single small collec-

tion that cannot be relocated in the field.

Vouchers: Bolus 5420 (BOL, K, LD, NBG, PRE, STE, W); Burchell

7773 (K, P, W); Oliver 8813 (BM , BOL, E, G, MO, MEL, NY, P, PRE,

S, STE, UPS, W); Oliver 9081 (E, MEL, PRE, STE); Schlechter 9639

(BM, BOL, E, G, K, MO, P, PRE, STE, W); Schlechter 10265 (BM,

E, K, MO, P, PRE, S, STE, W); Schlechter 10339 (BM, G, K, MO,

P, PRE, S, W); Sieber 165 (G, G-DC, K, LD, M, MO, P, PRE, S, W).

200 other collections studied.

3. Erica ericoides (L.) E.G.H. Oliver, comb. nov.

Blaeria ericoides L., Species plantarum 1: 112 (1753); L.: 331 (1771);

Willd.: 629 (1798); Thunb.: 7 (1802); Wendl.: 73 (1808); Ait.: 248 (1810);

Roem. & Schult.: 168 (1818); Bartl.: 649 (1832); Klotzsch: 663 (1833);

G. Don: 804 (1834); Klotzsch: 222 (1838); Benth.: 698 (1839); Rach:

788 (1853); N.E. Br.: 325 (1905); Aim & Fries: 233 (1924); Salter: 658

(1950); Bond & Goldblatt: 239 (1984); Oliver: 145 (1987). E. blaeria

Thunb.: 72 (1794); Thunb.: 358 (1823). E. dumosa Salisb.: 296 (1796);

Salisb.: 341 (1802). Lectotype: Hermann s.n. in Hermann Herbarium

vol. 4 fol. 61 (BM, STE, photo!), selected by Oliver (in press B).

Blaeria affinis N.E. Br. : 325 (1905). Type: Caledon Div., mountains

near Vogel Gat, near the mouth of the Klein River, 1500 ft., Schlechter

10418 (K, holo. ! ; BM!, BOL!, P!, PRE!).

This species, the type of Linnaeus’ genus Blaeria, is

very common in dry rocky areas on flats or in the moun-

tains from the Cape Peninsula eastwards along the coast

to just beyond Stanford (Figure 3). In some areas it forms

dominant stands which are very evident due to the strong

honeylike scent emitted by the flowers.

E. ericoides is a very distinct species which is often con-

fused with species from some of the minor genera, namely

Sympieza labialis (Salisb.) Druce, Simocheilus purpu-

reus (Berg.) Druce and Syndesmanthus articulatus (L.)

Klotzsch, all of which have similar looking heads of pale

pink tubular flowers with exserted dark brown anthers. The

Erica can be easily identified by its 4-celled multi-ovuled

ovary which produces a dehiscent capsular fruit.

Aim & Fries correctly placed Brown’s B. affinis in

synonymy under this species. The species was based on

the smaller flowerheads and finer branches of a single

collection from the Hermanus area. The numerous

collections of E. ericoides show a range of variation which

includes this slighter form.

E. ericoides is allied to E. russakiana and to a lesser

extent E. barbigeroides (q.v.).

The typification of E. ericoides is being formalised in

conjunction with R.F. Barrie (BM) for the programme on

the typification of Linnaean taxa (Oliver in press B).

Vouchers: Schlechter 7554 (BM, E, G, K, MO, P, PRE, STE, UPS,

W, Z); Schlechter 10401 (BM, BOL, E, G, K, MO, P, PRE, S, STE,

FIGURE 3. — The known distribution of Erica ericoides.

4

Bothalia 23,1 (1993)

FIGURE 4. — The known distribution of Erica fuscescens, # ; E. klotzschii , O; and E. multiflexuosa , ▲.

UPS, W); Sieber 172 (G-DC, K, LD, M, MO, P, S, STE, W). 170 other

collections examined.

4. Erica fuscescens (Klotzsch) E.G.H. Oliver, comb.

nov.

Blaeria fuscescens Klotzsch: 657 (1833); Benth.: 697 (1839); N.E. Br.:

321 (1905) pro parte excl. synonym Erica sagittate Klotzsch ex Benth.;

Aim & Fries: 239 (1924). Blairia fuscescens Dietr.: 443 (1839). Type:

Cape of Good Hope, Mundt & Maire s.n. (Bf, holo.; E!, G!, K!, P!,

W!). Lectotype chosen here: Mundt & Maire s.n. [det. Klotzsch] (P!).

This species is common on the southern slopes of the

Outeniqua and Tzitzikama Mountains in the southern Cape

(Figure 4). It has also been recorded from the Van Stadens

Mountains where it is sympatric with the closely related

E. sagittata (see below), but does not, as far as I am aware,

grow together with it. The numerous white flowers with

exserted black anthers make this a very striking plant. The

plants are bushy and fairly large, up to 1.5 m tall, whereas

in E. sagittata they are much smaller and sparser with

more crowded flowers due to the different inflorescence

type.

Vouchers: Burchell 5910 (BOL, K, P, UPS); Fourcade 831 (BOL, K,

STE, Z); Galpin 3717 (BOL, K, PRE, SAM); Mundt & Maire s.n. (E,

G, K, P, W) [ex B, det. Klotzsch]; Oliver 9250 (PRE, STE). 65 other

collections examined.

5. Erica klotzschii (Aim & Fries) E.G.H. Oliver,

comb. nov.

Blaeria klotzschii Aim & Fries: 237 (1924). Type: Cape of Good Hope

Lichtenstein s.n. (B|, holo.; S fragm.!). B. pusilla Klotzsch: 659 (1833)

nom. illegit. non Blaeria pusilla L.: 39 (1767) [= Simocheilus purpureus

(Berg.) Druce], non Erica pusilla Thunb.: 70 (1794), non Erica pusilla

Salisb. : 374 (1802); Benth.: 698 (1839); N.E. Br.: 322 (1905). Lecto-

type chosen here: Lichtenstein s.n. (S!).

E. klotzschii was rather anomalous in the genus Blaeria

as it had no close relatives and possessed only a superficial

resemblance to E. longimontana [— B. coccinea ]. The

flowers are very small and are borne at the ends of most

lateral branches on the plant thus producing plants almost

completely covered by pale pink to dull cream flowers.

The anthers are relatively large, exserted and slightly

versatile. This fact coupled with the dull colour of the

corolla, the small size of the flower and the far-exserted

cyathiform stigma strongly suggest wind pollination in

this species. This was substantiated in populations near

Swellendam in which plants gave off clouds of pollen when

disturbed early in the morning.

The species occurs on dry shaley hills and slopes at the

base of mountains from Elim to Swellendam and eastwards

as far as Albertinia often associated with Renosterveld

vegetation (Figure 4).

Vouchers: Bolus 8461 (BM, BOL, K, UPS); Oliver 4297 (BM, PRE,

STE); Zeyher 3331 (BOL, GRA, K, P, PRE, S, SAM, W, Z). 40 other

collections examined.

6. Erica longimontana E.G.H. Oliver, nom. nov.

Blaeria coccinea Klotzsch: 657 (1833), non Erica coccinea L. (1753);

Benth.: 697 (1839); N.E. Br.: 321 (1905); Aim & Fries: 236 (1924). Type:

Cape of Good Hope, Mundt & Maire s.n. (Bf, holo.; K!— BOL, fragm.!,

S! , W!). Lectotype chosen here: Mundt & Maire s.n. [ex B & det.

Klotzsch] (K!).

B. fastigiata Benth. : 697 (1839), non E. fastigiata L. (1771); N.E. Br. :

321 (1905). Type: Cape Colony, Burchell 7331 (K, holo.!; S!).

This species is common along the southern slopes of

the Langeberg from Swellendam to the western Outeni-

qua Mountains at the Robinson’s Pass (Figure 5) where

it is often dominant in seepage zones, hence the new name

chosen for the species.

Aim & Fries were correct in reducing Bentham’s B.

fastigiata to synonymy. This latter species was separated

off on the grounds of having only finely puberulous parts.

This form is found in the Swellendam area were it co-exists

with the form with longer hairs.

Vouchers: Oliver 9106 (BM, E, K, MEL, MO, NY, P, PRE, STE);

Schlechter 2055 (BM, BOL, G, K, PRE, S, UPS). 42 other collections

examined.

7. Erica multiflexuosa E.G.H. Oliver, nom. nov.

Blaeria flexuosa Benth.: 698 (1839), non E. flexuosa Andr. (1798);

N.E. Br.: 324 (1905). Type: at Steenbrass River, prov. Stellenbosch, Niven

s.n. (Herb Lambert; G-DC!). Lectotype chosen here: Niven 6 (Herb.

Lambert in K!).

B. purpurea Aim & Fries: 239 (1924) pro parte.

This species is confined to a small area on the lower

northern slopes of the Kogelberg complex around the

Steenbras Dam (Figure 4) where it grows in open sandy

areas between low restiad clumps. It forms low rounded

Bothalia 23,1 (1993)

5

but sparse shrublets with very intertwined branches which

are sparsely leafy. The flowers are a dull cream colour

and, hanging downwards, are rather inconspicuous.

E. multiflexuosa belongs to the E. equisetifolia complex

but can be distinguished by its intertwined branches, its

obovoid, dull cream, pendent flowers and included

muticous yellow-brown anthers. Like other species of the

complex it can have opposite leaves on some branches.

Brown (1905) was somewhat confused about this species

because he cited Niven 6 &. 7, both from the Steenbrass

River, under Blaeria flava and Niven 6 also under Blaeria

flexuosa.

Specimens examined

CAPE. — 3418 (Cape Town): Kogelberg Forest Reserve, E foothills of

Spinnekopsnes Range, 854 m, (-BB), 2-05-1970, Boucher 1285 (K, STE);

Steenbrass Reservoir valley, (-BB), 14-12-1933, Galpin 12405 (K, PRE);

Steenbrass area, (— BB), 1-05-1948, Levyns 8879 (BOL); ibid., Levyns

U533 (BOL); ibid., 7-12-1926, Middlemost 122 (BOL); ibid., Niven 6

(G-DC, K— BOL fragm.); ibid., Niven 7 (K); NE end of Kogelberg range,

450 m, (— BB), 20-03-1983, Oliver 8824 (PRE, STE); Steenbrass, Wol-

vem Kloof, (— BB), 28-02-1931, Stokoe 2563 (BOL, PRE, STE); Steen-

brass plateau & vlakte, (-BB), 18-02-1921, Stokoe 8917 (BOL); ibid.,

1931, Stokoe 8889 (BOL, PRE); near Steenbrass Reservoir, (-BB),

14-12-1933, Salter 4207 (BM, K, SAM).

8. Erica russakiana E.G.H. Oliver, nom. nov.

Blaeria kraussiana Klotzsch ex Walpers: 728 (1843) non E. kraussiana

Klotzsch ex Walpers: 728 (1843); Klotzsch ex Walpers: 824 (1844); N.E.

Br.: 326 (1905); Aim & Fries: 235 (1924); Oliver: 267 (1984). Type:

Babylon’s Tower, Hemel en Aarde, Krauss 973 (Bf, holo.; BOL!, K!,

M!, S!, UPS!, W!, Z!). Lectotype chosen here: Krauss 973 (K!-BOL

fragm.!).

Acrostemon concinnus N.E. Br. : 351 (1905). Type: Swartberg, Caledon,

Bodkin sub Bolus 9228 (BOL, holo.!; K!).

It is surprising that Brown overlooked the similarity

between his species and that of Klotzsch which he included

in his revision, especially seeing that his taxon was the

only one in Acrostemon with a 4-locular ovary.

The species, of which only four collections exist, is

confined to the northern slopes of the Klein River

Mountains north of Hermanus with one collection from

the Swartberg at Caledon (Figure 5). It forms a sparsely

branched low compact shrublet up to 300 mm tall

which becomes erect and more sparse when very old. The

flowers are superficially similar to those of the common

E. ericoides which occurs in the same area but forms a

large, woody spreading shrub up to 1.0 x 1.5 m. The other

species with similar heads of flowers, E. barbigeroides,

from the same area but near the southern base of the moun-

tains, forms a sparsely branched erect shrublet with much

larger and more numerous flowers per head.

The name chosen above continues to commemorate the

original collector, Ferdinand Krauss, but in anagrammatic

form.

Specimens examined

CAPE.— 3419 (Caledon): Swartberg, Caledon, (-AB), 01-1901, Bodkin

sub Bolus 9228 (BOL, K); Babylon’s Tower near Hemel-en-Aarde,

183-244 m, (- AC/AD), 08-1838, Krauss 973 (BOL, K, M, MO, S, UPS,

W, Z); Klein River Mtns above Diepgat, 610 m, (—AD), 1-02-1971, Oliver

3246 (PRE, STE); ibid., 22-02-1985, Oliver 8688 (STE).

9. Erica sagittata Klotzsch ex Benth. in DC., Prodro-

mus 6: 681 (1839). Type: in Vanstaaden mountains, Dr£ge

s.n., (Bf, holo. —BOL, sketch). Lectotype chosen here:

Vanstaadesberg [Van Stadensberg], 1000 ft., 27-12-1829,

Drege 7725 (P!).

Blaeria sagittata (Klotzsch ex Benth.) Aim & Fries: 238 (1924).

B. grandis N.E. Br.: 320 (1905). Syntypes: Van Stadens Mountains,

Zeyher 718 (K!, BOL fragm.!); ibid. West sub MacOwan 3119 (BOL!,

PRE!, SAM!).

B. juscescens sensu N.E. Br. : 320 (1905) quoad specm. Drege s.n.

Brown (1905) noted that Bolus had examined the type

of Erica sagittata in the Berlin Herbarium and had

informed him that despite its poor quality ‘there can be

no doubt of its identity with Blaeria juscescens Klotzsch’.

Bolus himself made a sketch of the type and some dissec-

tions of the flower and noted in his herbarium that it

contained three branches 12—14 inches long on one sheet

and only one imperfect flower in a capsule and that the

ticket labelled it as Drege 7725 from Van Stadensberg.

Bentham noted the type as ‘defloratum cum fl. unico

delapso’. If the specimen had possessed more flowers

Bolus would have seen that the axillary inflorescence on

an absolute brachyblast differed from the terminal

3-flowered inflorescence on a leafy lateral branchlet in

Blaeria juscescens and that the two species cannot be

confused.

The rather sparse material of Drege 7725 in Paris, con-

sisting of five young plants with two flowers in place on

FIGURE 5.— The known distribution of Erica longimontana , •; E. russakiana, A ; and E. sagittata , O.

6

Bothalia 23,1 (1993)

one plant, clearly matches the sketches of Bolus. It is

undoubtedly from the same collection as the lost Berlin

material.

This species is very localized in the Van Stadens Moun-

tains west of Port Elizabeth (Figure 5) where it is sympatric

with E. fuscescens. E. sagittata can be distinguished from

the latter species on a number of characters. Apart from

the type of inflorescence mentioned above, it has larger

flowers with corolla 6 mm long, not less than 4 mm, leaves

open-backed and 1—2 mm broad, not sulcate and 0.5— 1.0

mm long, and branches glabrous and distincdy ridged, not

puberulous and rounded.

E. sagittata is remarkable in being very similar to Erica

camea L. from Europe which also has flowers borne on

absolute brachyblasts and exserted dark stamens. The two

species could easily be mistaken for one another if the

origin of the material were not known. This is undoubtedly

a case of convergent evolution.

Specimens examined

CAPE.— 3324 (Steytlerville): Elandsrivierberg area in Otterford Forest

Reserve, 762 m, (-DB), 9-09-1973, Oliver 4455 (MO, PRE, STE). 3325

(Port Elizabeth): Blueberg, Loerie Plantation, 22-09-1934, (— CC), Dix

31 (BOL, GRA, K, PRE, STE); Van Stadens Gorge, Witteklip, (-CC),

09-1981, Muller s.n. (STE); Van Stadens Mtns, 305 m, (-CC/CD),

27-12-1829, Drige 7725 (P); ibid. , 09-1909, West 477 (BOL, SAM); ibid.,

West sub MacOwan 3119 (BOL PRE); ibid.. West sub MacOwan 3111

(BOL); ibid., Zeyher 718 (BOL, K); ibid., Zeyher 3266 (SAM).

INSUFFICIENTLY KNOWN SPECIES

Blaeria barbigera (Salisb.) G. Don: 805 (1834). TyPe:

Hottentots Holland, Masson s.n. (?).

Erica barbigera Salisb.: 341 (1802).

Salisbury’s type does not appear to exist, as no specimen

referrable to this species collected either by Masson or

labelled as ‘Herb. Salisb.’ nor one determined by Salisbury,

has been found in Kew or the British Museum, where all

the types of Salisbury’s species of Cape Ericaceae are

known to be housed. Salisbury’s description is insufficient

to be of use to ascertain the true identity of his species.

No mention was made of either the distinct capitate heads

of flowers (10—24) or of the glandular nature of the leaves

and calyx. His description could refer to a number of

species in Blaeria or even some other genera in the

subfamily. Without an authentic specimen it is not possible

to tie down Salisbury’s concept ‘ Erica barbigera'.

Bentham (1839) referred Salisbury’s species to Blaeria

revoluta Bartl. without having seen a type since he did

not cite a Masson collection. This was taken up by most

subsequent authors with Brown citing the synonymy as

‘sec. Benth.’.

EXCLUDED SPECIES

With the description of nunierous species under the

genus Blaeria during the late 1700’s and early 1800’s there

are many combinations under Blaeria which are no longer

in use for species removed to other genera. A full list of

these is given by Aim & Fries (1924: 262, 263) based on

the taxonomy of the family in Flora capensis (Brown

1905). With a revision of the family in southern Africa

currently in progress, many changes to the taxonomy and

therefore nomenclature, will be necessary. For this reason

a listing of all the excluded species is not published at this

stage as new combinations would have to be created of

which many would soon be redundant.

However, one species not covered by the listing of

Aim & Fries is: Blaeria muirii L. Guthrie: 179 (1924) =

Thoracosperma galpinii N.E. Br.: 330 (1905).

REFERENCES

AITON, W.T. 1810. Hortus kewensis edn 2, vol. 1. London.

ALM, C.G. & FRIES, T.C.E. 1924. Monographic der Gattung Blaeria

L. Acta Horti Bergiani 8: 223—267.

ANDREWS, H.C. 1794—1830. Coloured engravings of heaths I— IV.

London.

BARTLING, F. 1832. Plantae Ecklonianae, Ericeae. Linnaea 7: 627—652.

BENTHAM, G. 1839. Ericaceae. In A.P. De Candolle, Prodromus sys-

tematis naturalis regni vegetabilis 7: 580 —733. Paris.

BERGIUS, P.J. 1767. Descriptiones plantarum ex Capite Bonae Spei.

Salvius, Stockholm.

BOLUS, H. 1894. Contributions to the flora of South Africa. Journal

of Botany 1894: 239.

BOND, P. & GOLDBLATT, P. 1984. Plants of the Cape Flora, a descrip-

tive catalogue. Journal of South African Botany, Suppl. Vol. 13.

BROWN, N.E. 1905. Ericaceae. In W.T. Thiselton-Dyer, Flora capen-

sis 4: 315-336.

DAVIDSON, L.E. 1985. A change in status for Erica subverticillaris

var. revoluta from the eastern Transvaal. South African Journal

of Botany 51: 71—73.

DIETRICH, D.N.F. 1839. Synopsis plantarum. Vol. 1. Weimar.

DON, G. 1834. A general system of gardening and botany. Vol. 3.

London.

GUTHRIE, L. 1924. Novitates afficanae. Annals of the Bolus Herbarium

3: 179.

GUTHRIE, L. 1928. Novitates africanae. Annals of the Bolus Herbarium

4: 21.

GUTHRIE, L. & BOLUS, H. 1904. Erica. In W.T. Thistelton-Dyer,

Flora capensis 4: 4-315. Reeve, London.

KLOTZSCH, J.F. 1833. Ericearum a cel. Adelberto de Chamisso descrip-

tarum. Linnaea 8: 655-669.

KLOTZSCH, J.F. 1838. Ericearum genera et species. Linnaea 12:

211-247.

LINNAEUS, C. 1753. Species plantarum, edn 1. Stockholm.

LINNAEUS, C. 1767. Mantissa plantarum: 65 —66. Stockholm.

LINNAEUS, C. 1771. Mantissa plantarum alterum: 229—236.

Stockholm.

LINNAEUS, C. ftl. 1782. Supplementum plantarum. Braunschweig.

OLIVER, E.G.H. 1984. Studies in the Ericoideae. IV. New species and

some taxonomic and nomenclatural changes in the Cape Flora

Region. South African Journal of Botany 3: 267—284.

OLIVER, E.G.H. 1987a. Studies in the Ericoideae. VII. The placing

of the genus Philippia into synonymy under Erica-, the southern

African species. South African Journal of Botany 53: 455 —458.

OLIVER, E.G.H. 1987b. Ericaceae. In G.E. Gibbs Russell et al.. List

of species of southern African plants. Memoirs of the Botanical

Survey of South Africa No. 56: 140—147.

OLIVER, E.G.H. 1988. Studies in the Ericoideae (Ericaceae). VI. The

generic relationship between Erica and Philippia in southern

Africa. Bothalia 18: 1—10.

OLIVER, E.G.H. 1989. The Ericoideae and the southern African

heathers. Botanical Journal of the Linnean Society 101: 319—327.

OLIVER, E.G.H. 1993. Studies in the Ericoideae (Ericaceae). XHI. Three

new species of Erica from the southwestern Cape. Bothalia 23:

9-14.

OLIVER, E.G.H. in press A. Studies in the Ericoideae (Ericaceae). XI.

The generic relationship between Erica and Blaeria. Kew Bulletin.

OLIVER, E.G.H. in press B. In C. Jarvis & F.R. Barrie, Linnaean typifi-

cation. Regnum Vegetabile.

RACH, L. 1853. Die Ericaceen der Thunbeig’schen Sammlung. Linnaea

26: 788.

ROEMER, J.J. & SCHULTES, J. A. 1818. Systema vegetabilium. Cottae,

Stuttgart.

Bothalia 23,1 (1993)

7

SALISBURY, R. A. 1796. Prodromus stirpium in horto ad Chapel Allerton

vigentium. London.

SALISBURY, R. 1802. Species of Erica. Transactions of the Linnean

Society 6: 316—388.

SALTER, T.M. 1950. Ericaceae. In R. Adamson & T.M. Salter, Flora

of the Cape Peninsula. Juta, Cape Town.

THUNBERG, C.R 1794. Prodromus plantarum capensium Part 1: 1—83.

Uppsala.

THUNBERG, C.P. 1802. Dissertatio botanicae de Blaeria (P. Elmstedt):

1-12. Uppsala.

THUNBERG, C.P. 1823. Flora capensis ed. J.A. Schultes: 344-373.

Cotta, Stuttgart.

WALPERS, G.G. 1843. Ericaceae. Repertorium Botanices Systematicae

2 : 728.

WALPERS, G.G. 1844. Pflanzen des Cap- und Natal-Landes, gesam-

melt und zusammengestellt von Dr Ferdinand Krauss. Flora 48:

824.

WENDLAND, J.C. 1808. Collectio plantarum Vol. 1, fa sc. 1. Hannover.

WILLDENOW, C.L. 1798. Caroli a Linne species plantarum Vol. 1.

Berlin.

Bothalia 23,1: 9-14 (1993)

Studies in the Ericoideae (Ericaceae). XIII. Three new species of

Erica from the southwestern Cape

i E.G.H. OLIVER*

Keywords: Cape Province, Erica, Ericoideae, new species, taxonomy

ABSTRACT

With the recent inclusion of the genus Blaeria under Erica, three new species, formerly regarded as ‘incertae’ in that

genus, are now described: Erica chiroptera E.G.H. Oliver from the Kogelberg Reserve, Erica hermani E.G.H. Oliver and

Erica ioniana E.G.H. Oliver, both highly localised endemics from Hermanus.

UITTREKSEL

Met die onlangse insluiting van die genus Blaeria by Erica, word drie nuwe spesies wat voorheen onder die ‘incertae’

van daardie genus behoort het, nou beskryf: Erica chiroptera E.G.H. Oliver van die Kogelberg Reservaat, Erica hermani

E.G.H. Oliver en Erica ioniana E.G.H. Oliver, albei uiters beperkte endemiese spesies van Hermanus.

INTRODUCTION

The genus Blaeria L. was recognised for over two

centuries as separate from Erica L. based on the single

character difference of four as opposed to eight stamens

in the latter. In a recent paper (Oliver in press) it is shown

that there is an overlap between the two genera and that

there is a clear case for regarding Blaeria as polyphyletic.

As a result the genus Blaeria is being reduced to synonymy

under Erica (Oliver 1993).

Material which had been placed in the ‘incertae’ under

Blaeria and which clearly constituted several new taxa,

has remained undescribed because of the problem with

the delimitation of Blaeria. These taxa are now described

in this paper.

Erica chiroptera E.G.H. Oliver, sp. nov. in genere

distincta propter antheras quatuor inclusas latissimas

dorsiventraliter complanatas, capite aliquorum vespertilio-

num (Chiropteridae) similes, flores albos ad roseos;

affinitate E. equisetifoliae Salisb. sed facie antherarum

exsertarum differt.

Fruticulus erectus ad 500 mm altus. Rami erecti vel

flexuosi glabri trigoni, cortice griseo. Folia 3-nata, 1.5— 3.5

x 0.5— 1.0 mm, erecta appressa, angusta, sulcata glabra

ciliata vel eciliata, breve petiolata. Flores 3 — 6[9] ad

extremis ramorum et ramulorum lateralium; pedicellus

1. 5-3.0 mm longus glaber; bractea submediana 0.5— 1.3

mm longa lineari-oblonga glabra ciliata; bracteolae

bracteae similes minores. Calyx 4-lobatus, cyathiformis

0.8— 1.5 mm longus, glaber; lobi deltoidei, sulcati, sparse

ciliati pihs parvis et glandibus sessilibus. Corolla 4-lobata

2.7-3.6 x 2.0— 2.4 mm quadrangularis cyathiformis base

porcis calyce alternantibus glabra pustulate- rugosa; lobi

late rotundati irregulariter crenulati ad subfunbriati.

Stamina 4 inclusa; filamenta 0.9— 1.6 mm longa recta

glabra; anthera 0.9— 1.4 x 0.8— 1.4 mm erecta terminalia

* Stellenbosch Herbarium, National Botanical Institute, RO. Box 471,

Stellenbosch 7599.

MS. received: 1992-06-09.

late obcuneata dorsiventraliter complanata marginibus

strigulosis, thecis patentibus poro rotundato 'u longitu-

dine thecae append iculatis, cristis late obcuneatis decur-

rentibus. Ovarium 4-loculare 0.7— 1.0 X 0.7— 1.0 mm

distincte quadrangulare late ellipsoideum ad obovoideum

glabrum, nectario reducto vel absenti, ovulis 2 in quoque

loculo, ex placenta apicali pendulis, laterahter compressis;

stylus exsertus 2.0— 2.5 mm longus teres base expansus

glaber; stigma simplex. Fructus late ellipsoideus ad

obovoideus, septis longitudine 2>3 connatis, valvis ellip-

soideis, 1.5 x 1.2 mm, obtusis; semina ellipsoidea,

subreticulata. Figura 1.

TYPE. — 3418 (Simonstown): Cape, Caledon District,

Kogelberg Forest Reserve, neck at head of the Rooi Els

valley near Wynand Louwsbos, 366 m, (-BD), 11

November 1970, Oliver 3097 (STE, holo.; BM, BOL, E,

K, MEL, MO, NY, P, PRE, S, isotypes).

Erect, open or compact shrublet up to 500 mm tall.

Branches erect or wiry and flexuose, glabrous, with slight

infrafoliar ridges and trigonous when young, bark grey

flaking irregularly or mainly below leaves, becoming red-

brown. Leaves 3-nate, erect and appressed, subimbricate,

narrowly ovate to narrowly elliptic to linear-elliptic,

1.5— 3.5 x 0.5— 1.0 mm, subobtuse to acute, markedly

convex abaxially, flat or slightly concave adaxially, sulcate,

glabrous, with or without short fine cilia and marginal

sessile glands, strigulose at apex; petiole 0.3— 0.5 mm long,

glabrous. Flowers 3— 6[9] at ends of main and short sub-

terminal lateral branchlets or brachyblasts, pendent;

pedicel 1.5— 3.0 mm long, terete, glabrous, red; bract

submedian, linear-oblong, 0.5— 1.3 mm long, glabrous,

ciliate, sometimes with marginal sessile glands also, low-

er bracts in an inflorescence larger and sometimes sul-

cate, others not sulcate; bracteoles 2 just above bract and

like bract, 0.3— 0.7 mm long. Calyx 4-lobed to half its

length, cyathiform, 0.8— 1.5 mm long, glabrous; lobes nar-

rowly to broadly deltoid, clasping base of corolla, sulcate

in upper half, green, sparsely ciliate with short hairs and

sessile glands. Corolla 4-lobed, quadrangular cyathiform,

2.7— 3.6 x 2.0— 2.4 mm, dirty white to very pale pink,

4-ridged at base with ridges alternating with calyx lobes,

10

Bothalia 23,1 (1993)

FIGURE 1. — Erica chiroptera: A, flowering branch, x 2; B, close-up of branch with leaves; C, flower; D, androecium; E, anther, side, front

and back views; F, gynoecium; G, ovary, longitudinal section; B— G, X 25. All drawn from the type, Oliver 3097 (STE).

glabrous, pustulate-rugose; lobes erect to slightly spread-

ing, broadly rounded, '/4 the length of corolla, irregularly

crenulate to subfimbriate. Stamens 4 included; filaments

0.9— 1.6 mm long, straight, erect, slightly broadening

downwards, glabrous; anthers erect, placed just above

ovary, terminal, broadly obcuneate, with appendages

0.9— 1.4 X 0.8— 1.4 mm, dorsiventrally flattened, strigulose

on edges; thecae spreading in a broadly V-shaped arrange-

ment, 0.5— 0.9 mm long; pore rounded, 'u the length of

theca; appendages broadly obcuneate, decurrent for just

more than half their length. Ovary 4-locular, distinctly

quadrangular, broadly ellipsoid to obovoid, 0.7— 1.0 x

0.7— 1.0 mm, glabrous, nectaries very reduced or absent,

septa joined for half their length from base; ovules 2 per

locule, pendulous one above other from apical placenta,

laterally compressed; style exserted, 2.0— 2.5 mm long,

terete, enlarging slightly below stigma, occasionally bent

at apex, with enlarged basal portion forming a cap on ovary

apex, glabrous; stigma simple. Fruit broadly ellipsoid to

obovoid, septa joined for 2/3 of their length, valves

broadly elliptic, obtuse, 1.5 x 1.2 mm; seeds ellipsoid,

light brown, subreticulate. Figure 1.

E. chiroptera is a very distinct species on account of

its very broad, dorsiventrally flattened anthers which are

reminiscent of the heads of some bats (Order Chiroptera),

hence the name. It is, however, related to the extremely

variable species complex which is now referred to as E.

equisetifolia (Oliver 1993) and which occurs in the same

region. This latter species forms smaller, more compact

rounded shrublets which can, however, become taller and

erect when old. Its flowers are pink in colour, sometimes

deeply so, and have exserted anthers which are narrow and

elongate with small awns.

The species is confined to the Kogelberg Reserve

between Grabouw and Betty’s Bay (Figure 2) where it

grows in sandy quartzitic ground with short restiads.

Specimens examined

CAPE. — 3418 (Simonstown): Kogelberg Reserve, lower slopes south

of Klein Palmiet River, southwest of Somersfontein, 244 m, (— BB),

27-01-1987, Oliver 8999 (BM, BOL, G, K, NY, PRE, STE, W); Palmiet

River near Elgin, (— BB), 12-1941, Stokoe 6247 sub SAM 55121 (NBG,

PRE, SAM); Kogelbeig Reserve, Somersfontein boundary, 275 m, (— BD),

19-11-1969, Boucher 874 (PRE, STE); Kogelberg Reserve, Louws River

road, 335 m, (— BD), 11-03-1970, Boucher U79 (PRE, STE); Betty’s Bay,

plateau behind Cascades, 450 m, (— BD), 10-02-1956, Levyns 10456

(BOL); Kogelberg Reserve, head of Rooi Els valley near Wynand

FIGURE 2.— Known distribution of Erica chiroptera, •; and E.

hermani, O.

Bothalia 23,1 (1993)

11

Louwsbos, 366 m, (— BD), 11-03-1970, Oliver 3097 (BM, BOL, E, K,

MEL, MO, NY, P, PRE, S, STE). 3419 (Caledon): Arieskraal, (— AA),

30-12-1944, Leighton 915 (BOL); near Elgin, between Grabouw and the

Paardeberg, (-AA), 12-1947, Stokoe sub SAM 62387 (NBG).

Erica hermani E.G.H. Oliver, sp. nov. ex planitie

oraria Hermani, distincta propter folia sessilia appressa,

flores roseos campanulatos, stamina 8, antheras atras

strigosas exsertas mox deciduas; affinitate E. equisetifoliae

Salisb. ex eodem loco sed floribus tubulosis, staminibus

4 rare 5-8 (alteris locis) fuscis ad fulvis non deciduis

differt.

Fruticulus erectus ad 500 mm altus. Rami erecti, trigoni,

infra folia porcati, glabri roseo-brunnescentes, rami veteres

cortice frustris griseis. Folia 3-nata sessilia, appressa

subimbricata, 1.5— 2.8 mm longa, elliptica ad oblongo-

elliptica, acuta, glabra apice sparse strigulosa glandibus

sessilibus marginalibus. Flores 3— 9-nati terminales

pendentes; pedicellus 2.0— 2.8 mm longus, teretus, glaber;

bractea mediana, 0.7— 1.0 mm longa, naviculata, glabra

esulcata, breve ciliata pilis vel glandis sessilibus; bracteo-

lae subapproximatae bracteae similes sed breviores. Calyx

4-lobatus, obconicus ad cyathiformis, 1.2— 1.4 mm longus,

'u — !/3 connatus, base emarginatus; lobi ovato-deltoidei

ad late deltoidei, longe sulcati, ciliati glandis sessilibus

atrosanguineis. Corolla 4-lobata 2.4— 3.5 x 2.5— 3.0 mm,

obconica ad campanulata, interdum late campanulata et

parum quadrangularis, pustulata, pallido- ad atrorosea;

lobi longitudine '/4 — */3 corollae partes aequantes, late

deltoidei et obtusi, erecti ad plus minusve patentes,

irregulariter crenato-fimbriati. Stamina 8 exserta; filamen-

ta linearia ± 3 mm longa, sub anthera angustiora, glabra;

antherae bene exsertae, prope base dorsaliter affixae, post

anthesin mox deciduae, thecis 1.3— 1.5 mm longis, erectis,

appressis, suboblongo-ovatis, apice prognathis, obtusis.

marginibus strigosis ad base barbatis, nigris. Ovarium

4-loculare 0.6-0.8 x 0.6 -0.8 mm, quadrangulare, late

ellipsoideum, glabrum, base nectariis parvis; ovula 7-8

in quoque loculo, ex placenta apical i pendentes; stylus

exsertus, ± 5.4 mm longus, anguste cylindraceus basaliter

amplificatus, glaber; stigma obconica. Fructus late ellip-

soideus, septis longitudine 2/3 connatis, valvis 1.2 x 1.0

mm, late ellipsoideis, truncatis ad emarginatis; semina

complanata ellipsoidea, subreticulata, hepatica. Figura 3.

TYPE. — 3419 (Caledon): Cape, Hermanus, Fernkloof,

sandy lower slopes, 45 m, (-AD), 18 February 1984,

Oliver 8412 (STE, holo.; BM, BOL, E, G, K, MEL, MO,

NY, P, PRE, S, W, isotypes).

Sparse, erect shrub to 500 mm tall. Branches very fine,

erect, mostly leafy, trigonous when young with inffafoliar

ridges, glabrous, becoming round and reddish brown when

old with grey inffafoliar flakes of bark. Leaves 3-nate,

sessile, appressed, subimbricate, 1.5 -2.8 mm long, ellip-

tic to oblong-elliptic, acute, glabrous, sparsely strigulose

at apex, with a few sessile marginal glands when young.

Flowers 3-9-nate at ends of branches, pendent; pedicel

2.0— 2.8 mm long, terete, glabrous; bract median, 0.7— 1.0

mm long, naviculate, glabrous, without sulca, shortly

ciliate with hairs and/or sessile glands; bracteoles sub-

approximate, like bract but slightly shorter. Calyx 4-lobed,

obconical to cyathiform, 1.2— 1.4 mm long, joined for

*/3 — its length, emarginate at base, green to reddish,

lobes ovate-deltoid to broadly deltoid, long sulcate, edged

with sessile dark red glands. Corolla 4-lobed, 2.4— 3.5 x

2.5— 3.0 mm, obconical to campanulate sometimes broadly

so and somewhat quadrangular, pustulate, light to dark

cerise-pink; lobes ’/4 — ‘/3 the length of corolla, broadly

deltoid and obtuse, erect to slightly spreading, irregular-

ly crenate-fimbriate. Stamens 8 exserted; filaments linear,

FIGURE 3.— Erica hermani: A, flowering branch, x 2; B, close-up of branch with leaves; C, flower; D, anther, side front and back views;

E, ovary; F, ovary, longitudinal section; B-F, x 25. All drawn from the type, Oliver 8412 (STE).

12

Bothalia 23,1 (1993)

± 3 mm long, narrowed below anther, glabrous; anthers

well exserted, dorsally attached near base and soon

deciduous after anthesis, thecae 1.3— 1.5 mm long, erect,

appressed, suboblong-ovate, bent forwards at apex, obtuse,

strigose edged to bearded at base, black. Ovary 4-locular,

0.6— 0.8 x 0.6— 0.8 mm, quadrangular, broadly ellipsoid,

glabrous, with very small nectaries at base; ovules 7-8

per locule, pendent from an apical placenta; style exserted,

± 5.4 mm long, narrowly cylindrical with enlarged base

forming a cap to ovary, glabrous; stigma obconical. Fruit

broadly ellipsoid, septa joined for 2/3 their length, valves

1.2 x 1.0 mm, broadly elliptic, obtuse to emarginate; seeds

flattened ellipsoid, subreticulate, brown. Figure 3.

Erica hermani is allied to the variable and widespread

E. equisetifolia, formerly Blaeria equisetifolia (Salisb.)

G. Don and now including Blaeria dumosa Wendl. (Oliver

1993). It differs in having constantly eight stamens with

black, strigose, muticous anthers which are shed very soon

after anthesis; the flowers are also open-campanulate as

opposed to tubular. The shedding of the anthers leaves the

black-tipped filaments clearly visible in the mouth of the

corolla. The anthers of E. equisetifolia are variable in

shape being mostly long and narrow with decurrent awns

in the plants from the Hermanus area. Plants from the

inland mountains, formerly referred to Blaeria campanula-

ta Benth. and now placed under E. equisetifolia (Oliver

1993), have rather similar anthers to E. hermani but with

decurrent awns.

This species is very restricted, occurring only on the

sandy coastal flats near the entrance to the Fernkloof

Nature Reserve at Hermanus, hence the specific epithet,

(Figure 2) where it is locally common in remnant patches

of fynbos. There it co-exists with plants of E. equisetifolia

which looks remarkably similar, but is easily distinguished

on close examination of the flowers.

In the population of the above two species a few plants

with slightly different looking flowers were noted and on

close examination were found to be intermediates between

the two. The possession of nonviable pollen strongly

suggested a hybrid origin for these plants.

Specimens examined

E. hermani

CAPE. — 3419 (Caledon): Hermanus, Fernkloof, (—AD), 05-1986,

Drewe s.n. (STE); ibid., 45 m, 18-02-1984, Oliver 8412 (BM, BOL, E,

G, K, MEL, MO, NY, P, PRE, S, STE, W); ibid., 300 m, 12-01-1981,

Williams 3109 (BOL, PRE, STE); Mossel River flats, (—AD), Williams

sub Baker 3043 (STE).

E. equisetifolia [= Blaeria equisetifolia (Salisb.) G. Don]

CAPE.— 3419 (Caledon): Hermanus, sandy lower slopes at Fernkloof,

45 m, (—AD), 18-02-1984, Oliver 4111 (K, MEL, MO, NY, PRE, STE).

E. hermani x E. equisetifolia

CAPE.— 3419 (Caledon): Hermanus, sandy lower slopes at Fernkloof,

45 m, (—AD), 18-02-1984, Oliver 8413 (PRE, STE).

Erica ioniana E.G.H. Oliver, sp. nov. ex montibus

Hermani in genere distincta propter habitum parvum com-

pactum, flores campanulatos cremeos viscidos in capitibus

pendulis occultos, stamina straminea 4 interdum 5;

affinitatibus E. ericoidis (L.) E.G.H. Oliver et E.

russakianae E.G.H. Oliver sed inflorescentibus erectis,

floribus roseis non viscidis, staminibus fuscis.

Fruticulus compactus ad 150 mm altus. Rami glabri vel

sparse pubescentes ad villosi, sterigmatis infrafoliaceis,

rami veteres irregulariter porcati cicatricibus griseis. Folia

4-nata, 3.5— 5.0 mm longa oblongo-elliptica sparse villosa,

breviter glandulociliata; petiolus appressus, 1.2— 1.5 mm

longus, ciliatus pilis longis et glandibus sanguineis sub-

sessilibus. Flores 12—20 capitis compactis ad extremis

ramorum, dependentes, sub planta occultae; pedicellus

1.3— 1.7 mm longus, sparse villosus; bractea subapproxi-

mata 1.8— 2.5 mm longa, linearis ad lineari-spathulata,

foliacea in dimidio supemo sulcata, sparse villosa,

glandulis sessilibus vel subsessilibus ciliata; bracteolae

approximatae, 1.3— 2.0 mm longae, lineares ad filiformes

aliter bracteae similes. Calyx 4-lobatus, 1.4— 1.8 mm

longus; lobi subliberi anguste oblongo-lanceolati sparse

villosi glandulis subsessilibus sanguineis in marginibus et

in pagina adaxiali apicem versus, sulcatus. Corolla

4-lobata, 2.5 mm longa, campanulata quadrangularis,

glabra, viscida, cremea ad rosea; lobi 1 mm longi expansi

ad recurvi. Stamina 4(5); filamenta 3 mm longa, linearia,

apice latiore flexo; antherae inclusae ad submanifestae,

bipartitae, thecis 0.8 mm longis subellipsoideis, glabris,

super ovario impendentibus; porum longitudine '/3 thecae

partes aequans; pollen in tetradis. Ovarium 4-loculare, 0.6

x 0.6 mm, late ovoideum, 4-lobatum, plus minusve

emarginatum, glabrum, base nectariis grandibus; stylus

2 mm longus, exsertus; stigma simplex. Fructus globosus

vel late ovoideus, valvis 1.0 x 0.8-1.0 mm late ellipticis,

obtusis ad emarginatis; semina ovoideo-ellipsoidea,

reticulata, fusca. Figura 4.

TYPE. — 3319 (Caledon): Cape, Hermanus, Mossel

Neck at SW end of Vogelgat Nature Reserve, 490 m,

(-AD), 22 February 1985, Oliver 8684 (STE, holo. ; BOL,

K, MO, PRE isotypes).

Low compact shrublet to 150 mm tall with slightly

spreading branches. Branches glabrous or sparsely pubes-

cent to villous when young, with slight infrafoliar ridges

when young, bark with slight infrafoliar flakes otherwise

flaking irregularly when old. Leaves 4-nate, 3.5— 5.0 x

3.0 mm long, oblong-elliptic, sparsely villous all over and

sometimes shortly gland-ciliate, becoming strigose; petiole

appressed, 1.2— 1.5 mm long, ciliate with long hairs and

red subsessile glands. Flowers 12—20 in compact heads

at ends of branches, feeing downwards, hidden under plants

and not visible; pedicel 1.3— 1.7 mm long, sparsely vil-

lous, red; bract subapproximate, 1.8— 2.5 mm long, shorter

than calyx, linear to linear-spathulate, foliaceous and sul-

cate in upper half, sparsely villous, edged with sessile to

subsessile red glands; bracteoles approximate, 1. 3-2.0

mm long, shorter than calyx, linear to filiform, otherwise

similar to bract. Calyx 4-lobed, 1.4— 1.8 mm long; lobes

almost free, narrowly oblong-lanceolate, just shorter than

corolla tube, sparsely villous, with subsessile red glands

on edges and adaxially towards apex, sulcate, hairy,

green turning red. Corolla 4-lobed, 2.5 mm long,

campanulate, 4-angled, glabrous, viscid, cream to pale

pink; lobes 1 mm long, spreading to recurved. Stamens

4(5); filaments 3 mm long, linear with a broader bent apex;

anthers included to just manifest, bipartite, pale yellow

FIGURE 4. — Erica ioniana : A, flowering branch, x 2; B, whorl of leaves; C, flower; D, sepal; E, anther, side front and back views; F, gynoecium;

G, mature ovary; B— G, x 25. All drawn from the type, Oliver 8684 (STE).

brown; thecae 0.8 mm long, slightly ellipsoid, glabrous,

bent forwards over ovary; pore V3 the length of theca;

pollen in tetrads. Ovary 4-locular, 0.6 x 0.6 mm, broadly

ovoid, 4-lobed, slightly emarginate, glabrous, with well-

developed nectaries at base; style 2 mm long, exserted;

stigma simple. Fruit globose or broadly ovoid, valves 1.0

x 0.8— 1.0 mm, broadly elliptic, obtuse to emarginate;

seeds ovoid-ellipsoid, reticulate, dark brown. Figure 4.

This distinct species was brought to my attention by Dr

Ion Williams of Hermanus (alter whom the species is

named) who is undertaking surveys of his Vogelgat Reserve

in the Klein River Mountains above Hermanus. He

assumed the plant to be, from the description, Blaeria

affinis N.E. Br. which is known only from the type

collection made in Vogelgat by Schlechter. This latter

species was reduced to synonymy under E. ericoides

(L.) E.G.H. Oliver (= B. ericoides L.) by Aim & Fries

(1924). I have found that Schlechter ’s collection is just a

depauperate form of the common species.

A visit to the area revealed that Williams’ populations

were a distinct new species with an unusual habit. The

plants are low, compact and rounded with the flowers

arranged in pendent heads such that they are not easily

noticeable when the plants are in full bloom. The pale

colour of the corolla and the long hairs of the calyces in

the heads of flowers also help to make the flowers less

visible. In these respects the species is unlike any other

in the area. The other species there with capitate inflores-

cences, namely E. ericoides, E. russakiana and E.

barbigeroides E.G.H. Oliver, have the branches spreading

or erect and different, dark brown, basally winged or

appendiculate, exserted anthers. The anthers in E. ioniana

are pale yellowish brown, muticous, included and are

narrowed in at the base. They are very similar to those

found in E. multiflexuosa E.G.H. Oliver, a restricted

endemic near Steenbras Dam.

The species is restricted to level sandy places in and

near the Vogelgat Reserve, Hermanus, (Figure 5) where

three populations have so far been found. The unusual

habit strongly suggests some form of pollination by

creeping insects with the flowers facing downwards only

a short distance from the ground.

Specimens examined

CAPE. — 3419 (Caledon): Hermanus, Mossel Neck in Vogelgat

Reserve, 490 m, (—AD), 22-02-1985, Oliver 8684 (BOL, K, MO, PRE,

STE); ibid., 480 m, (—AD), 9-02-1985, Williams 3580 (NBG, PRE);

Caledon District, north side of Klein River Mtns above Diepgat, 594

m, (-AD), 22-02-1985, Oliver 8688 (BM, E, K, MEL, NY, PRE, STE);

Hermanus, Femkloof, Mt Pustulata, 610 m, (—AD), 26-02-1984, Williams

3531 (HER, NBG, PRE, P, S).

REFERENCES

ALM, C.G. & FRIES, T.C.E. 1924. Monographic der Gattung Blaeria

L. Acta Horti Bergiani 8: 223—267.

FIGURE 5.— Known distribution of Erica ioniana.

Bothalia 23,1 (1993)

OLIVER, E.G.H. in press. Studies in the Ericoideae (Ericaceae). XI. placing of the genus Blaeria into synonymy under Erica-, nomen-

The generic relationship between Erica and Blaeria. Kew Bulletin. clatural changes for the southern African region. Bothalia 23:

OLIVER, E.G.H. 1993. Studies in the Ericoideae (Ericaceae). XII. The

1-7.

-

. >.

Bothalia 23,1: 15-23 (1993)

The hepatics, Symphyogyna podophylla and Pallavicinia lyellii (Palla-

viciniaceae) in southern Africa

S.M. PEROLD*

Keywords: Hepaticae, Pallaviciniaceae, Pallavicinia lyellii, southern Africa, Symphyogyna podophylla, systematics

ABSTRACT

A detailed description of Symphyogyna podophylla, illustrated with photographs of the thalli and spores, is presented.

It is compared to S. hymenophyllum, with which it may be conspecific. The subcosmopolitan Pallavicinia lyellii, also occurring

in southern Africa, is described and illustrated with photographs.

UITTREKSEL

’n Uitvoerige beskrywing van Symphyogyna podophylla, gefllustreer met foto’s van die tallusse en spore, word gegee. Dit

word vergelyk met S. hymenophyllum waaraan dit gelyksoortig mag wees. Die subkosmopolitiese Pallavicinia lyellii, wat

ook in suidelike Afrika voorkom, word beskryf en met foto’s gefllustreer.

INTRODUCTION

Symphyogyna podophylla is locally quite widespread but

of rarer occurrence in southern Africa than S. brasiliensis,

the other species recorded for the region. The latter was

recently redescribed and illustrated with photographs

(Perold 1992) in order to draw attention to its southern

African synonyms Pallavicinia capensis, Symphyogyna

valida and S. lehmanniana (Grolle 1980). Sporulating

material of S. podophylla held at PRE is rare, Koekemoer

991 (which was recently collected), being only the fourth

specimen of this species which has spores.

Amell (1963), Vanden Berghen (1965), Grolle (1979) and

Grolle & Piippo (1986) regard S. hymenophyllum (Hook.)

Nees & Mont, as conspecific with S. podophylla, but

this has been questioned, since there appear to be some

differences in the spore ornamentation, as seen on SEM

micrographs and in the chemistry of plants from New

Zealand (E.O. Campbell pers. comm.). This description

of 5. podophylla, illustrated with photographs, is given

in the hope that it may help to answer the question whether

S. hymenophyllum is conspecific with S. podophylla or

not.

The subcosmopolitan Pallavicinia lyellii of the same

family Pallaviciniaceae Migula, but different subfamily

Pallavicinioideae (Migula) Grolle, is also redescribed and

illustrated with photographs. Differences between the

genera Symphyogyna and Pallavicinia are briefly discussed

in Perold (1992) and under Pallavicinia lyellii below.

Symphyogyna podophylla (Thunb.) Mont. & Nees

in Gottsche, Lindenberg & Nees, Synopsis hepaticarum:

481 (1846); Amell: 107 (1963); Vanden Berghen: 157

(1965). Type: Cape, ‘Promontorium Bonae Spei’, leg.

Thunberg (25945 UPS, holo.!; S, STR, W, iso.).

Jungermannia podophylla Thunb.: 174 (1800); Thunb.: 738 (1823).

* National Botanical Institute, Private Bag X101, Pretoria 0001.

MS. received: 1992-07-29.

S. rigida Steph.: 112 (1914); Steph.: 69 (1917). Syntypes: Rwanda,

Rugege-Wald: ca. 1 900 m, leg. Mecklenburg 842', Vulkan-gebiet:

Ninagongo ca. 3 500 m, leg. Mecklenburg 2012 (?G). [Syn. fide Vanden

Berghen: 159 (1965)].

S. rhizobola (Schwaegr.) Nees & Mont.: 68 (1836). Jungermannia

rhizobola Schwaegr.: 31 (1814). Type: Reunion (W, iso.). [Syn. fide

Grolle: 267 (1979)]. See Amell: 109 (1963).

S. harvevana Taylor: 408 (1846). Type: Cape of Good Hope, leg.: W.H.

Harvey (FH, holo.; NY, S, W, iso.). [Syn. fide Sim: 33 (1926)].

Terricolous, on damp soil; thallus, mostly erect,

dendroid shoots in loose mats or tufts, green to olive-green,

medium-sized; proximal branches, after initial dichotomy,

single, ± 4 mm long, then a further 2 or 3 times dichoto-

mously furcate, with 7 or 8(-12) terminal branches, linear

to narrowly ovate, up to 11 mm long, 1.6 -2.0 mm wide,

125 thick over ventrally slightly bulging costa, lacking

rhizoids, but with central, brown conducting strand clearly

visible and forking at dichotomies; apex rarely progres-

sively narrowed with a reversion to stipe-like condition

and arching down, mostly entire or slightly to deeply

notched, bearing 2- celled slime papillae, margins of wings

dentate, lacking slime papillae, plane, not undulating or

hardly so, bilaterally expanded from ascending wingless

stipe (Figure 1H), 4—18 mm long, in transverse section

275 x 500 gm, arising from horizontally creeping and

much branched, cylindrical brown rhizome, with nume-

rous smooth ventral rhizoids, ± 12.5 pm wide. Wings

generally unistratose, but medianly bistratose and grading

into flatfish costa (Figure IK), laterally with marginal

teeth, remote or closer together, occasionally very blunt,

usually with 2(or 3) forwardly directed cells, diverging

at an angle of ± 45°, basal cell 62.5 x 37.5 pm (rarely

with 2 adjacent cells), top cell bluntly conical, 50 x 30

pm\ marginal cells ± rectangular to polygonal

(40.0— )67.5— 90.0 x 27.5—37.5 pm, 30-35 pm thick in

transverse section, their walls and those of l(or 2) rows

of inframarginal cells generally somewhat thicker (Figure

U), coloured pinkish; laminal cells arching toward

margins, polygonal, up to 82.5 x 55.0 pm; epidermal cells

along costa narrowly rectangular or long-hexagonal,

75.0-125.0 x 37.5 pm, occasionally bearing 2-celled slime

papillae above; chloroplasts numerous, ± 5 pm wide,

16

Bothalia 23,1 (1993)

FIGURE 1. — Symphyogyna podophylla, plants and androecia. A, male branches in situ; B, branches with rows of antheridial scales along costa,

C, some antheridial scales above costa, much enlarged; D, E, excised antheridial scales; F, apex of lobe; G, laminal cells with numerous

round chloroplasts and few spindle-shaped oil bodies; H, transverse section of stipe; I, transverse section of costa and central conducting

strand, much enlarged; J, transverse section of lateral and thicker-walled marginal cells of lobe; K, transverse section of lobe; L, toothed

margin of lobe from above. A— F, H— L, Koekemoer 991; G, Koekemoer 994. A, x 4; B, x 5; C, x 70; D, E x 100; F, H, L, x 125;

G, x 500; I, J, X 250; K, x 50. A— L, LM photographs.

FIGURE 2. — Symphyogyna podophylla, female plants. A, B, female branches with capsules above bifurcation ot costa; (J, part ot snoot calyptra

in transverse section; D, young seta in transverse section; E, involucral scale and cluster of archegonia; F, outer cells of capsule wall;

G, transverse section of capsule wall; H, spore and elater. A— G, Koekemoer 991. A, X 10; B, x 8; C, x 100; D, X 125; E, X 50;

F, G, x 250. A— G, LM photographs.

entirely filling cells, or clustered along cell walls, several

oil bodies also present, spindle-shaped (Figure 1G) and

+ 7.5 pm long, or round when viewed end-on. Costa with

central conducting strand, 37.5 x 50.0 pm, consisting

mostly of 12 small, ± 10 x 10 pm, brown, thick-walled,

angular cells, surrounded above and below by 3 rows of

larger, thin-walled parenchymatous cells (Figure II),

20.0-22.5 x 25.0-37.5 pm.

Dioicous. Androecia in 1 or 2 dorsal rows over the cos-

ta (Figure 1A— C); antheridia 200 x 175 pm, short-stalked,

each one individually covered and well hidden by irregu-

larly shaped, forwardly directed scale-like involucre

(Figure ID, E), 175 pm wide at base, up to 450 pm long,

irregularly incised at apex. Gynoecia generally 2 per

frond, dorsally situated above costa at bifurcation of

conducting strand (Figure 2A, B), containing several

archegonia and subtended by posteriorly inserted involucre

(Figure 2E), 675 pm wide at base, deeply laciniate to

filiform above, cells short- to long-rectangular, 45.0-125.0

x 71.5—41.5 pirn. Calyptra thickening and enlarging into

a fleshy shoot calyptra, ± 4 mm long and up to 8 cell

18

Bothalia 23,1 (1993)

rows or 250 /xm thick in transverse section (Figure 2C),

with several unfertilized archegonia remaining attached

near the top (Figure 2 A). Capsule cylindrical , 1800 x 850

pm, opening along several longitudinal valves, remaining

attached above, wall brown, 2 cell layers thick (Figure 2G),

outer cells elongate (Figure 2F), 62.5— 107.5 x 17.5 pm,

walls thickened, inner cells thin-walled. Seta erect when

young, 480 /im in diameter, with ± 36 cortical cells, ±

50.0 x 32.5 )tm, medullary cells slightly larger, 62.5 x

37.5 pm, angular, thin-walled (Figure 2D). Spores light

brown, ± globular, 20—25 pm in diameter, ornamenta-

tion nodular, with some very irregular, broad flattened

granular ridges (Figure 3A— C), meshes sometimes

distinct, but mostly not; proximal face with small round

area with compact, punctate ornamentation (Figure 3D,

E). Elaters brown, hardly tapering toward ends, 305—330

x 7.5 pm, 2-spiral (Figures 2H; 3F).

Symphyogyna podophylla is quite rarely collected in the

Transvaal and Natal, but fairly frequently in southwestern,

southern and eastern Cape. There are no records from the

Orange Free State (Figure 4). From other parts of Africa,

Amell (1960) has reported it from Ethiopia (= Abyssinia)

and Vanden Berghen (1965) has also been reported it from

Ethiopia (= Abyssinia) as well as from Tanzania and the

Congo Republic (Kivu Province) . Best (1990) records both

5. podophylla as well as its synonym, S. harveyana, from

Zimbabwe. Vana et al. (1979) report S. podophylla from

Zaire, Rwanda and Reunion; Bizot & Poes (1974) found

it to occur in Kenya on Mt Kenya and the Aberdare Moun-

tains in montane mossy forests and subalpine moorland;

Bizot et al. (1976) report it from Malawi. Some of these

specimens may, however, belong to Jensenia spinosa

(= Pallavicinia spinosa ), with which S. podophylla has

sometimes been confused (Grolle 1979).

DISCUSSION

In such a wide ranging species one would expect some

local variation. A Sim specimen, PRE-CH 1491, from

Table Mountain has bistratose wing margins, and medianly

there are four layers of cells. The determination was kindly

confirmed by Dr Grolle. Amell (1963) regarded the size

of the marginal teeth in S. podophylla as having no taxo-

nomic significance, nor the thickness of the walls in the

marginal cells. He also did not find distinct differences

between t;ie erect and procumbent forms, there being a

continued gradation between the two.

In a comparison of transverse sections taken at the

middle of the apical lobes of specimens of S. podophylla

and of S. hymenophyllum from Juan Fernandez, Arnell

(1956) stated that the thickness of the ‘nerves’ and the size

of the teeth in the margin of the thalli vary but little. Amell

(1963) also compared specimens of S. hymenophyllum from

Tristan da Cunha and Inaccessible Island as well as from

Reunion and from Kilimanjaro, with numerous specimens

of S. podophylla from the Cape (e.g. Thunberg’s collec-

tions) and could not find any real difference between them.

On the other hand, Gottsche et al. (1846) distinguished

S. podophylla from S. hymenophyllum by the ‘frondis

laciniis basi attenuatis subpetiolatis’ of the former.

In the present study, a photograph of the transverse

section of a lobe (Figure IK) was compared with the

drawing of a section, taken midway of a terminal

lobe, by Hassel de Menendez (1961a: fig. 10c) and the

resemblance is unmistakable. It may thus be that Arnell

(1963) was indeed correct in placing S. hymenophyllum

in synonymy under S. podophylla. Scott (1985) also recog-

nizes the Australian species as S. podophylla. Should

FIGURE 3. —Symphyogyna podophylla, spores and elaters. A, B, distal face; C, distal face much enlarged; D. part of proximal face with punctate

area; E, punctate area on proximal face, much enlarged; F, elaters. A, Koekemoer 991', B— E, Doidge 168', F, Hilner CH 1499. A, X 1835;

B, D, x 1530; C, E, x 2524; F, x 720.

Bothalia 23,1 (1993)

19

FIGURE 4. — Distribution of Symphyogyna podophylla, ▲; and

Pallavicinia lyellii, O, in southern Africa.

S. podophylla and S. hymenophyllum eventually conclu-

sively prove to be conspecific, then S. podophylla would

be circumsubantarctic in its range, since S. hymeno-

phyllum is reported to occur in South America, and

New Zealand is the type locality. Poes (1976), in discuss-

ing the presence of subantarctic bryophytes in tropical

Africa, remarks on the high number of subantarctic ele-

ments that are present and reckons that they may have

migrated northwards by means of mountain hopping.

Symphyogyna podophylla is a less common and smaller

plant than S. brasiliensis . It can be distinguished by: its

growth form which is generally flabellate from an erect

stalk arising from a creeping rhizome; the lobe margins

which are toothed and hardly undulate; it rarely fruits and

the spore ornamentation is characterised by broad, flat,

roughened, irregular ridges, sometimes forming ‘loops’,

and not by narrow, erect ridges as in S. brasiliensis.

Symphyogyna podophylla and S. brasiliensis are the only

two species of Symphyogyna occurring in southern Africa

and are placed in the subfamily Symphyogynoideae (Trev.)

Grolle.

Note: the collector of the type specimens of S. rigida is

stated to be Mildbraed by Stephani (1917) and Geissler &

Bischler (1990). However, Mildbraed was the publisher

of ‘Wissenschaftliche Ergebnisse der Deutschen Zentral-

Afrika-Expedition 1907—1908’; the leader of the

expedition, Adolf Friedrich, Duke of Mecklenburg, should

be cited as collector, as was done by Vanden Berghen

(1965) and now by me.

SPECIMENS EXAMINED

TRANSVAAL.— 2230 (Messina): Zoutpansberg, Piesanghoek, at

Waterfall, (— CC), M. Bosnian 206 (PRE); Piesanghoek, (-CC), M.

Bosnian 943 (PRE); Entabene, (-CC), M. Bosnian 1774 (PRE);

Entabene, (— CC), Bottomley 2865 (PRE); De Hoek State Forest,

Grootbosch Hiking Trail. ± 9.5 km from start, (— CC), Koekemoer 991

(PRE); Zoutpansberg, (— CC), Obermeyer 1941C ex herb. Tvl. Museum,

Obermeyer 3109 (PRE). 2329 (Petersburg): Haenertsburg. (— DD), H.A.

Wager CH 3808 (PRE). 2430 (Pilgrims Rest): Graskop, Fairyland,

(— DD), Glen 2982 (PRE); Hebron Mountain, in second cleft on plateau,

against vertical stream wall, (— DB), Vorster 1000 (PRE); Hebron

Mountain, northern slope, gallery forest along stream in plantation, on

stone and soil, (— DB), Vorster 1635 (PRE); Mariepskop, Blyde River

footpath, on wet earth bank along stream in forest, (— DB), Vorster 1516

(PRE). 2527 (Rustenburg): Magaliesberg, lower Tonquani KJoof, cliff

face, in dripping water, (-CA), H. Anderson 1249, 1250 (PRE); Magalies-

berg range. Crystal Waters, 13 km E of Rustenburg, in constant drip of

waters of sheer chasm, into a stream a few feet below, (— CA), Mogg

34818 (PRE). 2530 (Lydenburg), Kaapsche Hoop, (— DB), V.A. Wager

62 (PRE).

NATAL.— 2731 (Louwsburg): Zululand, Vryheid Dist., Ngoni Forest,

(- CD), Forester CH 1505, CH 1529, CH 1530 (PRE). 2828 (Bethlehem):

Mont-aux-Sources, (-DD), Doidge 168 (PRE). 2929 (Underberg):

Donnybrook, Gala Bush, (— DD), Doidge CH 13112 (PRE). 2930 (Pieter-

maritzburg): Edendale, Gordon Falls, (— CB), T.R. Sim CH 1495 (PRE);

Hilton Road, (- CB), T.R. Sim 8242 (PRE).

CAPE. — 3227 (Stutterheim): Hogsback, Victoria East, (— CA), Van

der Bijl 75 (PRE); Hogsback, in shade, streambank, (— CA), Young CH

1507 (PRE); Evelyn Valley, (-CB), T.R. Sim CH 1522 (PRE). 3318 (Cape

Town): Table Mountain, Disa Gorge, (—CD), S. Amell 1059, 1099, 3932

(PRE); Table Mountain, Window Gorge, (—CD), Bews CH 1517 (PRE);

Table Mountain, Skeleton Gorge, (—CD), Bews 8482 (PRE); top of Table

Mountain, (—CD), Bews 8516 (PRE); Table Mountain, Woodhead Tunnel

Gorge, (—CD), Bews 8519 (PRE); in spring at base of Lion’s Head,

(-CD), Bews CH 1490 (PRE); Table Mountain, (-CD) , Bolus CH 1496

(PRE); Table Mountain, Skeleton Ravine, (-CD), Bolus CH 1521 (PRE);

Table Mountain, (-CD), Michell CH /4<S9(PRE); eastern slopes of saddle

between Table Mountain and Devil’s Peak, on wet and partly shaded

rocks, (-CD), Pillans 3548 (PRE); Devil’s Peak, The Grottos, (—CD),

T.R. Sim CH 1492 (PRE); Table Mountain, Window Gorge waterfall,

(-CD), T.R. Sim CH 1493 (PRE); Table Mountain, Slongoli, (-CD),

T.R. Sim CH 1494 (PRE); Table Mountain, Woodhead tunnel intake,

(—CD), T.R. Sim CH 1506 (PRE); Cape Town, Platteklip stream, (—CD),

T.R. Sim CH 1511 (PRE); Newlands ravine, (—CD), T.R. Sim CH 1527,

CH 1528 (PRE); Table Mountain, (-CD), H.A. Wager 4 (PRE). Without

locality, H.A. Wager 3828 (PRE); Stellenbosch, Paradise Ravine, grow-

ing in dense masses, (— DD), (label in Duthie’s handwriting), ex Herb.,

Sim CH 1510 (PRE); Stellenbosch Mountain, above Brandwacht, lower

slopes facing west, wet streamside, (— DD), Oliver 9028 (PRE). 3319

(Worcester): Tulbagh Dist., Sneeugat Valley, (— AA), Thome CH 2877

(PRE); Groot Drakenstein Mountains, (— CC), Primos CH 1504 (PRE).

3320 (Montagu): Tradouw Pass, S of Barrydale near waterfall in forest-

ed kloof, (-DC), Magill 6174 (PRE). 3321 (Ladismith): Seven Weeks

Poort, (—AD), Thome CH 1543 (PRE). 3322 (Oudtshoom): Meiring-

spoort, in deep shade in cracks of vertical rock wall, almost under water-

fall, (— BC), Jacobsen 2253 (PRE); Montagu Pass, wet rock face near

Stinkhoutdraai, (—CD), S.M. Perold 1547 (PRE); The Wilderness, Ge-

orge, (—CD), Taylor CH 1519 (PRE); Saasveld, Groeneweide, (—DC),

Koekemoer 994 (PRE). 3326 (Fort Beaufort): near Grahamstown,

Paradise Kloof, Coldstream, (— BC), Hilner CH 1499 (PRE). 3227

(Stutterheim): Hogsback, (— CA), Van der Bijl CH 1497 (PRE); Hogsback,

in shade, streambank, (— CA), Young CH 1507 (PRE); Evelyn Valley,

(— CB), T.R. Sim CH 1522 (PRE). 3418 (Simonstown): kloof near Chaplin

Point, (— AB), T.R. Sim CH 1501 (PRE); Kogelberg, near Gordon’s Bay,

(-BB), Mogg CH 939 (PRE); Hottentot Hollands Mts, (— BB), Thome

CH 3096 (PRE). 3419 (Caledon): Riviera Kloof, Herman us Dist., (—AC),

Louwrens CH 2889 (PRE); Mossel River, (—AD), Potts 26 (PRE);

Oudebos, Zonder Einde, (— BB), Thome CH 3106 (PRE).

Pallavicinia lyellii (Hook.) Car ruth, in Journal of

Botany, British and Foreign 3: 302 (1865) ( Pallavicinius)\

Sim: 32 (1926); Muller: 519 (1951-1958); Hassel: 264

(1961b); Hodgson: 223 (1968); Grolle & Piippo: 60 (1986).

Jungermannia lyellii Hook.: tab. 77 (1816). Type: England, Hamp-

shire, New Forest, ‘Cadman bog’, 6 May 1812, Lyell (BM, lecto. fide

Grolle & Piippo 1986).

Pallavicinia pilifera Steph.: 271 (1891); Steph.: 10(1900); Amell: 112

(1963); Vanden Berghen: 164 (1965). Type: W Africa. Sao Tome Island,

Quintas (G 12058) (G, holo.; G, M, S, W, syn.). Synonymy fide Vanden

Berghen: 150 (1972).

For detailed synonymy see Grolle & Piippo (1986).

Terricolous, growing on damp soil; thallus, prostrate

and creeping (Figure 6A), in crowded, overlying,

caespitose mats, dark green, mostly simple, occasionally

dichotomously branched, or ventrally from the midrib,

medium-sized to large, up to 60 x (2.0— )4. 0—6.5 mm,

320—350 gm thick over ventrally bulging costa from

20

Bothalia 23,1 (1993)

which, at intervals, arise dense reddish brown, smooth,

translucent rhizoids 12.5—20.0 /xm wide; central conduct-

ing strand brown, clearly visible from above, forking at

dichotomies; apex entire or with shallow notch, bearing

2-celled slime papillae (Figure 5F), also along margins

of wings, which are undulating, almost entire or with

remote teeth, expanded bilaterally from wingless stipe,

375 x 750 (im in transverse section. Wings unistratose.

bordered by a row of mostly long-rectangular or 5-sided

cells (27.5— )50.0— 62.5 x 27.5—45.0 /xm, marginal teeth

generally small and blunt (Figure 5D), only 1- or 2-celled,

25.0 x 37.5 /xm, toward apex often larger, up to 5-celled,

forwardly directed (Figure 5E); laminal cells polygonal,

some hexagonal, 57.5—87.5 x 27.5—45.0 /xm, in transverse

section 42.5 /xm thick, containing numerous chloro-

plasts, 5.0— 7.5 /xm, oil bodies green, nodular, 8—12 per cell

FIGURE 5 .—Pallavicinia lyellii , plants and androecia. A, male branch with androecia laterally situated above costa; B, male scales more enlarged;

C, part of male scale, excised; D, margin of lobe, with blunt tooth; E, long marginal tooth, near apex; F, notched apex with 2-celled slime

papillae, and long marginal teeth; G, laminal cells from above, containing chloroplasts and rod-like oil bodies; H, transverse section of

costa and central conducting strand, much enlarged; I, transverse section of costa and wing on one side only. A-C, Glen 2882: D-I,

Koekemoer 990. A, x 30; B, X 70; C, x 165; D, x 250; E, F, H, X 125; G, x 500; I, x 25.

Bothalia 23,1 (1993)

21

FIGURE 6 .—Pallavicinia lyellii, female plants. A, lobe with gynoecia along costa; B, involucre much enlarged; C, opened involucre spread out;

D, lobe with capsule still enclosed in pseudoperianth; E, calyptra; F, pseudoperianth; G, transverse section of capsule wall; H, outer cells

of capsule wall; I, transverse section of seta; J, elater and spore. A— C, Koekemoer 990 ; D— F, I, Glen 2882\ G, H, J, Amell 2084. A,

-x 5.5; B, x 55; C, x 70; D, x 11; E, F, x 30; G, H, J, x 250; I, x 80.

(Figure 5G), up to 7.5 /tm long, thin, others seen end-on

and round. Costa abruptly grading into wings, overlying

epidermal cells long-rectangular, 65 — 110 x 21—30 /tm,

sometimes with slime papillae, central conducting strand

50 x 80—125 /tm, consisting of about 32 small, ± 10 /tm

wide, thick-walled, angular brown cells, surrounded by

larger, 35—55 /tm wide, thin-walled parenchymatous cells,

3 rows above and 4 or 5 rows below (Figure 5H).

22

Bothalia 23,1 (1993)

Dioicous. Androecia in 2 parallel dorsal rows, one on

either side of costa (Figure 5A), each a flat band raised

shelflike ± 100 pm above lamina and ± 280 pm wide,

laterally at free margin, expanded into confluent scales,

directed transversely to long axis of shoot (Figure 5B),

210 x 200 pm, with laciniae (l-)2- or 3-celled (Figure 5C),

± 150 x 40 /im; antheridia globular, 170 p m wide, short-

stalked and individually placed, hidden by covering scale,

central area above midrib lacking antheridia and scales.

Gynoecia up to 4 or 5 per frond (Figure 6A), acropetally

arranged and dorsally situated at intervals along costa,

involucre 3—5 layers thick, cup-like, base + 900 pm high,

mouth densely fringed with ciliate laciniae up to 750 pm

long (Figure 6B, C), enclosing a group of 20—30 archego-

nia. Pseudoperianth 6 mm long, tubular (Figure 6F), apex

brownish, long-piliferous (Figure 6D), developing after

fertilization of an archegonium; calyptra (Figure 6E) bis-

tratose, initially enveloping capsule and seta, several un-

fertilized archegonia flattened against sides and at base,

old archegonial neck retained at tip. Capsule oblong-

cylindrical, 2.3— 3.7 mm long, opening along several

valves; wall yellow-brown, 2 cell layers thick (only 1 shown

in Figure 6G); outer cells elongate, 87.5—150.0 x

12.5—22.5 pm, lacking semi-annular thickenings (Figure

6H). Seta with slight swelling at foot, eventually up to 9

mm long, sinuate, 500 pm in diameter, cortical cells

darker, one cell deep, ± 35 x 35 pm, in 45 cell rows;

medullary cells ± 70, + 35 —60 (im wide (Figure 61).

Spores light brown, + globular, 17.5— 23.0 /xm in di-

ameter, ornamentation reticulate with larger areolae on dis-

tal face ± 5 pm wide, further subdivided into smaller

areolae by finer walls (Figure 7A— D); proximal face with

only smaller, irregular areolae (Figure 7C— E). Elaters

light brown, tapering toward ends, up to 470 pm long, 7.5

pm thick, bispiral (Figures 6J; 7F).

northern and central Transvaal, Natal, and southwestern,

southern and eastern Cape (Figure 4), but not many

collections have been made. It is probably more common,

but is undercollected.

The very large number of synonyms listed by Grolle &

Piippo (1986) are an indication of its variability, resulting

from its plasticity. Fertile plants are rare, with male plants

generally smaller and abruptly narrowing to the apex of

the lobe. The marginal teeth are quite variable: mostly

small and blunt, but sometimes up to 5 cells long. In

colour, plants are a deeper green than those of Sym-

phyogyna species and further differ by the androecial

arrangement, by the cup-like involucre, short calyptra, the

presence of a pseudoperianth and by the spore ornamen-

tation. The plants described and illustrated as Pallavinicia

lyellii by Sim (1926) are clearly those of Symphyogyna

brasiliensis.

SPECIMENS EXAMINED

TRANSVAAL. — 2330 (Tzaneen): De Hoek State Forest, Grootbosch

Hiking Trail, + 6.5 km from starting point, (— CC), Koekemoer 990

(PRE). 2430 (Pilgrim’s Rest): Graskop, Fairyland, occasionally on dry

stream bank, (— DD), Glen 2975 (PRE); Mariepskop, Blyde River

footpath, on sandstone slab in forest, (-DB), Vorster 1499 (PRE). 2527

(Rustenburg): Rustenburg Nature Reserve, Cederbergkloof near Utopia,

(— CA), Koekemoer 971 (PRE).

NATAL. — 2731 (Louwsburg): Louwsburg Dist., Ngome Forest,

Cetshwayo waterfall walk, (—CD), Glen 2882 (PRE). 2930 (Pietermaritz-

burg): Blinkwater, (— AB), J.M. Sim CH 1456 (PRE).

CAPE.— 3318 (Cape Town): Table Mountain, (—CD), Michell CH 1481

(PRE). 3322 (Oudtshoorn): George, Van Riebeeck Gardens, on damp

streambank, (—CD), S.M. Perolcl 927 ( PRE). 3326 (Grahamstown): Para-

dise Kloof, Cold spring, near Grahamstown, (— BC), Hilner CH 1478

(PRE). 3423 (Knysna): Knysna, BuffelstNek, streamside, (— AA), S.

Arnell 1515 (PRE); Knysna, Garden of Eden, (— AA), S. Arnell 2084

(PRE).

DISCUSSION

ACKNOWLEDGEMENTS

Pallavicinia lyellii is widely distributed and regarded

as subcosmopolitan. In southern Africa it is known from

I wish to express my gratitude to the following: Dr E.O.

Campbell and Dr R. Grolle for refereeing the manuscript

FIGURE 1. —Pallavicinia lyellii, spores and elaters. A, B, distal face; C, D, side view; E, proximal face; F, elater. A-F, S. Arnell 2084. A,

B, D, x 1850; C, E, x 1905; F, x 189.

Bothalia 23,1 (1993)

23

and for their helpful suggestions; to my colleagues at

NBI, Ms Koekemoer and Dr H.F. Glen, for collecting

specimens; to Mrs A. Romanowski, photographer, for

developing and printing the photographs (taken by the

author) and to Mrs J. Mulvenna, typist, for her valued

contribution.

REFERENCES

ARNELL, S. 1956. Hepaticae collected by O. Hedberg et al. on the East

African mountains. Arkiv for Botanik 3: 517—562.

ARNELL, S. 1960. Hepatics collected by Dr John Eriksson in Abyssinia

in 1958. Svensk Botanisk Tidskrift 54: 187—192.

ARNELL, S. 1963. Hepaticae of South Africa. Swedish Natural Science

Council, Stockholm.

BEST, E.B. 1990. The Bryophyta of Zimbabwe— an annotated check-

list. Kirkia 13: 293-318.

BIZOT, M. & POCS, T. 1974. East African Bryophytes I. Acta Academiae

Paedagog. Agriensis, Eger n. Ser. 12: 383—449.

BIZOT, M., DURY, M.N. & POCS, T. 1976. East African Bryophytes

11. Acta Botanica Academiae Scientiarum Hungaricae 22:

1-8.

CARRUTHERS, W. 1865. On the nomenclature of the British Hepaticae.

Journal of Botany, British and Foreign 3 : 297 —302.

GEISSLER, P. & BISCHLER, H. (Eds) 1990. Index hepaticarum. Vol.

12. Racemigemma to Zoopsis.

GOTTSCHE, C.M., LINDENBERG, J.B.G. & NEES AB ESENBECK,

C.G. 1844—1847. Synopsis hepaticarum.

GROLLE, R. 1979. Miscellanea hepaticologica 171—180. Journal of

Bryology 10: 263—272.

GROLLE, R. 1980. Miscellanea hepaticologia 201—210. Journal of

Bryology 11: 325—334.

GROLLE, R. & PIIPPO, S. 1986. Bryophyte flora of the Huon Peninsula,

Papua New Guinea. XVI. Acta Botanica Fennica 133: 59—

79.

HASSEL DE MENENDEZ, G.G. 1961a. Las Especies Argentinas del

Genero Symphyogyna. Boletin de la Sociedad Argentina de

Botanica IX: 233— 260.

HASSEL DE MENENDEZ, G.G. 1961b. Las Especies Sudamericanas

del Genero Pallavicinia. Boletin de la Sociedad Argentina de

Botdnica IX: 261-282.

HODGSON, E.A. 1968. New Zealand Hepaticae (Liverworts) XVIII.

A review of the New Zealand species of the genera Pallavicinia

and Symphyogyna. Transactions of the Royal Society of New

Zealand, Botany 3: 221—230.

HOOKER, W.J, 1816. British Jungermanniae. Longman.

MULLER, K. 1951—1958. Die Lebermoose Europas. In Rabenhorst,

Kryptogamenflora 6: 416—471.

NEES AB ESENBECK, C.G. & MONTAGNE, C. 1836. Jungerman-

nicarum herbarii Montagneani species. Annales des Sciences

naturelles, Bot. Sdr. 2,5: 52—72.

PEROLD, S.M. 1992. The occurrence in southern Africa of the hepatic,

Symphyogyna brasiliensis (Pallaviciniaceae). Bothalia 22: 53—58.

POCS, T. 1976. Correlation between the tropical African and Asian

bryophytes, 1. Journal of the Hattori Botanical Laboratory 41:

95-106.

SCHWAEGRICHEN, C.F. 1814. Historia muscorum hepaticorum pro-

dromus. Leipzig.

SCOTT, G.A.M. 1985. Southern Australian liverworts. Australian Flora

and Fauna Series No. 2.

SIM, T.R. 1926. The Bryophyta of South Africa. Transactions of the

Royal Society of South Africa 15: 1—475.

STEPHANI, F. 1891. Hepaticae africanae. Hedwigia 30: 265 —272.

STEPHANI, F. 1900. Species hepaticarum 1: 305—400.

STEPHANI, F. 1914. In Mildbraed, Wissenschaftliche Ergebnisse der

Deutschen Zentral Afrika-Expedition 1907-1908 Band 2 (Botanik):

1-718.

STEPHANI, F. 1917—1924. Species hepaticarum 6: 1—693.

TAYLOR, T. 1846. New Hepaticae . The London Journal of Botany 5 : 408.

THUNBERG, C.P. 1800. Prodromus plantarum capensium. Part 2:

85-191.

THUNBERG, C.P. 1823. Flora capensis.

VAfaA, J., POCS, T. & DE SLOOVER, J.L. 1979. Hepatiques

dAfrique tropicale. Lejeunia 98: 1—23.

VANDEN BERGHEN, C. 1965. Hepatiques recoltees par le Dr J.J.

Symoens dans la region peri-tanganyikaise. Bulletin Societe Royale

de botanique de Belgique, Bruxelles 98: 129—174.

VANDEN BERGHEN, C. 1972. Hepatiques et Anthocerotees. Resultats

scientifiques de Vexploration hydrobiologique du Bassin Lac

Bangweolo & Luapula 8: 1—202.

Bothalia 23,1: 25-47 (1993)

A biosystematic study of Pentameris (Arundineae, Poaceae)

N.P. BARKER*

Keywords: Arundineae, Arundinoideae, cladistics, conservation status, cytology, leaf anatomy, Pentameris, phylogeny, systematics

This paper is dedicated to the memory of Lucy K.A. Crook (nee Chippindall)

ABSTRACT

A biosystematic study of the endemic southwestern Cape grass genus Pentameris Beauv. is presented. Results of studies

on the macro- and micromorphology, leaf blade anatomy and cytology are discussed and illustrated . The results of a cladistic

study indicate that the genus is monophyletic, united by the synapomorphies of ovary and fruit characters. The conservation

status of the taxa in the genus is assessed, and conservation status codes allocated. A key to the taxa in the genus is presented,

and each species is described. Five new species, Pentameris glacialis N.P. Barker, P. hirtiglumis N.P. Barker, P. oreophila

N.P. Barker, P swart be rgensis N.P. Barker and P. uniflora N.P. Barker, and one new subspecies, P longiglumis (Nees)

Stapf subsp. gymnocolea N.P. Barker, are described and illustrated.

UITTREKSEL

’n Biosistematiese ondersoek van die endemiese Suidwes-Kaapse grasgenus Pentameris Beauv. word aangebied. Resultate

van ondersoeke op die makro- en mikromorfologie, blaarskyfanatomie en sitologie word bespreek en gefllustreer. Die resultate

van ’n kladistiese ondersoek dui daarop dat die genus monofileties is, verenig deur die afgeleide kenmerke van die vrugbeginsel

en vrug. Die bewaringstatus van die taksons in die genus word geraam en bewaringstatuskodes toegeken. ’n Sleutel tot die

taksons in die genus word aangebied, en elke spesie word beskryf. Vyf nuwe spesies, Pentameris glacialis N.P. Barker,

P. hirtiglumis N.P. Barker, P oreophila N.P. Barker, P. swartbergensis N.P. Barker en P. uniflora N.P. Barker, en een

nuwe subspesie, P. longiglumis (Nees) Stapf subsp. gymnocolea N.P. Barker, word beskryf en geillustreer.

CONTENTS

Introduction 25

Morphology 25

Leaf blade anatomy 29

Microhairs 29

Abaxial macrohairs 29

Adaxial macrohairs 30

Prickles 30

Cytology 31

Phylogeny 31

Choice of outgroup 31

Character distribution 32

Taxonomic implications 33

Conservation status 33

Taxonomy 33

Acknowledgements 44

References 46

INTRODUCTION

The genus Pentameris Beauv. occurs in the winter

rainfall region of the Cape Province, South Africa, where

it is restricted to soils derived from Table Mountain

Sandstone or the shale bands associated with this geology.

It may therefore be considered an endemic of the Cape

Flora (Goldblatt 1978). All known species of the genus

are perennial C3 plants.

The genus was erected by Palisot de Beauvois in 1812

on the basis of a specimen sent to him by Du Petit-Thouars.

No collection number or locality is provided in this

description. However, it is known that Du Petit-Thouars,

* National Botanical Institute, Private Bag X101, Pretoria 0001. Present

address: Botany Department, University of Cape Town, Private Bag, Ron-

debosch, Cape Town 7700.

MS. received: 1992-02-20.

a French traveller and botanist, obtained material from the

Cape during a visit in February 1793 (Gunn & Codd 1981).

The genus name is Greek, meaning ‘five parts’, probably

a reference to the apical region of the lemma which is

divided into what Palisot De Beauvois (1812) described as

four bristles (two of which may be more accurately

described as lemma lobes) and a central awn.

Only one species is mentioned under the generic

description: P. thuarii Beauv. (Palisot de Beauvois /.c).

This single species was subsequently placed in Dantho-

nia DC. by a number of early taxonomists (Desvaux 1831;

Nees 1841; Steudel 1855; Durand & Schinz 1895), whereas

others retained it in the genus Pentameris (Roemer &

Schultes 1817; Kunth 1833, 1835). Stapf (1897) expanded

the genus Pentameris to include four other taxa, all charac-

terised by fruit with a free pericarp. Chippindall (1955)

retained the genus in this format, but incorporated two

nomenclatural corrections published by Schweickerdt

(1938). Gibbs Russell et al. (1985) list these five taxa as

Pentameris dregeana Stapf, P longiglumis (Nees) Stapf,

P macrocalycina (Steud.) Schweick., P. obtusifolia (Hochst.)

Schweick. and P thuarii Beauv.

However, the unusual leaf blade anatomy (Ellis 1985d;

Barker 1990) and fruit morphology (Barker 1986, 1989,

1990) of P. obtusifolia places this species within the genus

Pseudopentameris Conert (Barker in prep.a). Pentameris

obtusifolia is therefore not further considered in this study.

MORPHOLOGY

The basal parts of all the species are woody, but the

woolly and/or swollen underground parts that occur

in certain species of Pentaschistis (Nees) Spach and

Merxmuellera Conert are absent. The species are generally

tufted, with stems sometimes branching, but generally only

26

Bothalia 23,1 (1993)

in the basal quarter. In older plants branching results in

a cushion or a bush-like growth form. These branched

structures are referred to here as ‘culms’, although they

do not possess typical culm anatomy. Instead, the anatomy

resembles that of a rhizome (H.P. Linder pers. comm.).

The term ‘aerial culm’ will be used here to denote the

annually produced, few-noded structure which terminates

in an inflorescence.

In P. thuarii, branching can occur at any node on the

culm, and some specimens comprise long, decumbent,

scandent or vertical culms with many shorter vegetative

and reproductive branches arising from the upper regions.

Such branching from cauline innovation shoots results in

an ‘evergreen’, perennial plant, which takes the form of

a divaricate herb. This branching pattern might allow the

plant to grow taller than species possessing only basal

innovation shoots (Linder & Ellis 1990a). Branching and

growth from cauline innovation shoots is also found in a

few species of Pentaschistis, Chaetobromus Nees and

Pseudopentameris . In the latter genus the branched

condition may approximate the extreme development found

in Pentameris thuarii.

The leaf structure of Pentameris is typical of the

Arundineae. The leaf sheaths are persistent, although the

leaf blades may not be. The older culms are thus frequently

covered in the remains of sheaths from previous years of

growth. These sheaths may be appressed to or free from

the culm. The sheath indumentum varies between the spe-

cies, ranging from glabrous to pubescent along the mar-

gins to uniformly pubescent (and sometimes woolly).

These sheath characters are also found in other arundi-

noid genera. The ligule is, as in many other arundinoid

genera, a fringe of hairs. The sheath mouth may be beard-

ed in some species, but this, too, is not unusual. Two spe-

cies, P thuarii and P longiglumis have fairly

well-developed auricles. In the former species these are

a characteristic purple or brown colour. The leaf blade

varies in its internal and epidermal anatomy. Many charac-

ters are shared, although not consistently so, with taxa of

Pentaschistis, and (to a lesser extent) Merxmuellera. Ellis

(1985c, 1986; Ellis & Linder 1992), on the basis of leaf

anatomical studies, is of the opinion that Pentameris is

closely allied to Pentaschistis.

The inflorescence is a panicle which varies from com-

pactly lanceolate to laxly globose. It must, however, be

noted that the panicles of all the species are open and

somewhat lax during the period of pollination. Before and

after this period, the panicles contract to a lanceolate

shape. Panicle shape is therefore difficult to describe as

it depends on the reproductive phase of the plant at the

time of observation or collection. This variation is also

found in numerous other arundinoid taxa, for example

Pentaschistis (Linder & Ellis 1990b). No spicate panicles

are known from this genus.

The spikelets are two-flowered (one exception; P.

uniflora), and the partially developed remains of a third

floret may sometimes be present, particularly in P.

macrocalycina. The two-flowered state is shared with

the vast majority of the species in Pentaschistis (Linder

& Ellis 1990b). When two florets are present, the basal

floret is sessile, the apical floret pedicellate. The florets

are otherwise morphologically identical, and both are

hermaphroditic and fertile, as in Pentaschistis.

The lemma is generally nine- to eleven-nerved. The

majority of the veins anastomose in the basal region of

the central awn and the two lateral bristles. The lemma

lobes on the outside of the bristles are variously adnate

to the bristle, ranging from free (in P. thuarii) to almost

completely adnate (in P. macrocalycina). The shape of this

lobe is also variable, being acuminate, acute or dentate.

Similar variation is found in Pentaschistis (Linder & Ellis

1990b). The central awn is geniculate in all species, the

basal portion is flattened but twisted and shorter than the

attenuating apical portion. The margin of both the basal

and apical regions of the awn are finely serrated. This awn

structure is shared with genera such as Pentaschistis,

Pseudopentameris, Merxmuellera and Chaetobromus as

well as the non-African genus Danthonia.

The palea is bicarinate, apically bilobed and pubescent

between the veins. It exceeds the lemma body in length,

but seldom exceeds the lemma lobes, a situation also found

in species of Pentaschistis, Pseudopentameris and Merx-

muellera.

The lodicules in certain arundinoid genera have been

examined and used for taxonomic purposes (Tomlinson

1985). The lodicules of Pentameris are generally glabrous

and cuneate, but differences have been noted in some

specimens of P. swartbergensis and P distichophylla. The

lodicules in these species are generally apically ciliolate,

and sometimes an arm-like extension arising from one of

the lateral margins is present. Ciliolate lodicules are also

known from species of Pseudopentameris, Merxmuellera

and Pentaschistis (Linder & Ellis 1990b).

All species of the genus have three anthers, which are

usually purple in colour. These vary in size in relation to

the size differences of the florets, but this has not been

incorporated as a taxonomically meaningful character.

The above morphological features can therefore not be

used to clearly differentiate Pentameris from several other

southern African arundinoid genera, notably Pentaschistis.

There are, however, two micromorphological features

which have, in the past, been used to distinguish Penta-

meris from all other southern African arundinoids. Both

features are characters of fruit and ovary. Firstly, the fruit

is an achene, and secondly, the fruit of all taxa possesses

a dense tuft of hairs at the apex of the ovary. These hairs

are retained in developing and mature fruit where they

become reluctantly deciduous.

Stapf (1897: 512) separated Pentameris from Pentaschistis

on the basis of fruit morphology: the crustaceous pericarp

and free seed. He described the structure of the ovary as

being ‘. . . so alike in the five species of this genus that it

is very probable that they agree in the peculiarities of the

ripe fruit which is known only in P. thuarii' and further

stated that ‘... there is in Pentaschistis no approach to

the characteristic structure of the ovary and the fruit of

Pentameris .’

This unusual fruit type, the achene, is described by

Clayton & Renvoize (1986) as a fruit with a thin, hard,

free pericarp attached to the ovule in the region of the

hilum. It is unusual in the tribe Arundineae, and is found

in only three of the approximately 40 arundinoid genera

worldwide. The fruits of the remaining arundinoid genera

are described as caryopses or as caryopses with a variably

separable pericarp (Clayton & Renvoize l.c.).

Bothalia 23,1 (1993)

27

FIGURE 1 .—Pentameris; A, B, SEM of fruit; C— F, SEM of leaf blade. A, P. hirtiglumis N.P. Barker, whole fruit, note colliculate surface features

and apical hairs. B, D, F, P. thuarii Beauv.: B, longitudinal section; D, furrow on adaxial surface; F, ad axial surface showing ribs over

vascular bundles and open furrows between ribs. Note abundance of conventional macrohairs and slightly inclined, sparsely distributed prickles.

C, P. distichophylla (Lehm.) Nees, adaxial microhair. E, P. oreophila N.P. Barker, magnified view of deflated apical cell of adaxial microhair.

Ac, small deflated apical cell; Ah, apical hairs; Be, long basal cell; E, embryo; Em, endosperm; H, hilum; Ma, macrohair, basal portion;

Mi, microhair; P, prickles; Pe, free crustaceous pericarp, seed within; S, stomata. Scale bars: A, 390 /im; B, 541 fan; C, 19 pm; D, 33

/im; E, 6 jim; F, 550 pm.

Scanning electron microscope (SEM) studies on the fruit

of Pentameris have shown the pericarp to be sculptured

(Barker 1986, 1989, 1990) as illustrated here in Figure 1A,

and partly or almost completely free from the seed coat

when viewed in section (Barker 1990) as shown here in

Figure IB. The colliculate surface sculpturing is also

unique among the fruit of the southern African genera of

the Arundineae (Barker in prep. b).

The second feature that characterises the genus is the

presence of hairs on the apical region of the ovary and

fruit. This was first observed by Palisot de Beauvois (1812),

who described the fruit as being crowned with stellate

hairs. Nees (1841) also noted this feature in his descrip-

tions of P. thuarii and the taxon now known as P

macrocalycina. Phillips (1931) considered these hairs to

be characteristic of the genus, using this feature in his

TABLE 1.— Salient anatomical characters differentiating the species of Pentameris. All measurements in micrometres

28

Bothalia 23,1 (1993)

primary vascular bundles; 3' = tertiary vascular bundles.

Bothalia 23,1 (1993)

29

key to the grass genera. Chippindall (1955) describes

Pentameris as having an ovary which is hairy on top, the

hairs being deciduous. This character was also used in the

key to the genera of the tribe Danthonieae. The nature of

these unusual structures has been clarified to a certain

extent by Barker (1986, 1989, 1990), but their ontogeny

and function is not known. These structures, visible on

the apex of the fruit (Figure 1A) appear to be unicellular,

and arise from the apex of the ovule around the base of

the styles.

LEAF BLADE ANATOMY

Leaf blade anatomy has contributed much to the tribal

and subfamilial classification of the grass family. However,

within the tribe Arundineae, the variation of observed

anatomical characters led Renvoize (1981) to comment that

the arundinoid genera could not be readily divided into

the tribal groups on the basis of their leaf anatomy, and

he concluded that other characters such as spikelet

morphology would have to be used to divide the subfamily

further.

The leaf blade anatomy of many of the southern African

arundinoid genera has been documented (Schweickerdt

1942; De Wet 1956; Conert & Tuerpe 1969; Ellis 1980;

1981; 1985a— d).

Ellis (1985a— d, 1986) examined many specimens of

the then five known and one undescribed species of

Pentameris. Additional anatomical information for the new

and undersampled species was obtained by sectioning a

small portion of leaf material obtained from herbarium

specimens. This material was heated in a soapy solution

prior to sectioning by means of a sledge microtome.

Sections thus obtained were 30 to 50 microns thick. The

sections were stained in safranin and fast green and

mounted in Euparol.

To complement the data obtained from the work of Ellis

(1985a-d, 1986) and additional sections, leaf material

from 95 of these previously studied specimens was

prepared from herbarium specimens, mounted on two-

sided tape, coated in gold-palladium and examined using

a ISI-SX-25 scanning electron microscope. The salient

anatomical features obtained from the SEM and light

microscope studies are presented in Table 1.

The abaxial surface is relatively uniform throughout the

genus. It is generally smooth, with ribs observed only

occasionally. No abaxial microhairs or stomata are present,

and the silica bodies are usually rounded, but are tall and

narrow in P thuarii (Ellis 1985b— d, 1986). Abaxial

macrohairs were occasionally present in specimens of

some taxa (Table 1). The adaxial epidermis is far better

endowed with micromorphological structures, including

prickles, microhairs and macrohairs. These epidermal

features are discussed in some detail below.

Microhairs

The genus Pentameris has a ‘festucoid’ type of leaf

epidermis and leaf anatomy (implying the absence of

abaxial microhairs), a poorly differentiated parenchyma

sheath and evenly distributed chlorenchyma sheath (De

Wet 1956). However, Renvoize (1986) places Pentameris

in the ‘core’ of the arundinoids, a group possessing micro-

hairs, and describes the lower epidermis of Pentameris and

Pseudopentameris as possessing long slender papillae

which occasionally bear the remains of a small thin-walled

apical cell. Clayton & Renvoize (1986) note that these

structures distinguish these genera (and certain others such

as Cortaderia Stapf and Centropodia Reichb. f.) from the

rest of the Arundineae.

These ‘papillae’, as described by Renvoize (1986) and

Clayton & Renvoize (1986), appear to differ from those

described by Ellis (1979), who considers papillae to be

protuberances of the cell wall rather than structures

consisting of separate cells. From observations discussed

below, it appears that Renvoize is using the term ‘papillae’

to describe what others, such as De Wet (1956) and Ellis

(1979), call microhairs.

In addition to their structure and appellation, the

recorded distribution of these microhairs is also controver-

sial. Renvoize (1986, discussed above) reported them from

the lower (abaxial) surface of specimens of Pentameris.

However, Ellis (1985a-d, 1986) has found no evidence

of abaxial microhairs on any of the specimens he has

examined. Microhairs were however found on the sides

and bottoms of the furrows of the adaxial surfaces of many

specimens in all the taxa observed (Ellis l.c.). My obser-

vations on material examined, using the SEM, corroborate

Ellis’s observations, and it is clear that there are no abaxial

microhairs present in any of the species of Pentameris.

As illustrated in Figure 1C, D and E, these microhairs

are indeed bicellular, with a minute, deflated apical cell

(shown at high magnification in Figure IE). The length

of the basal cell varies, but it is always very much longer

than the apical cell. Microhairs with this structure were

found on the adaxial surface of all specimens of all taxa

in Pentameris (Barker 1990), and were also reported from

certain species of Pentaschistis (Ellis & Linder 1992).

In addition to their distribution and morphology, varia-

tion in the size of the microhairs is taxonomically useful.

The microhairs were measured from a number of SEM

micrographs in order to determine whether or not there

was any measurable size difference. These varying sizes

are recorded in Table 1. Unfortunately, the limited sample

size does not allow a statistically meaningful compari-

son to be carried out on this potentially very valuable

taxonomic character.

The definite bicellular nature of these microhairs is

comparable to those reported and illustrated by

Amarasinghe & Watson (1988, 1989). However, microhairs

with such unequal cell sizes have not been previously

documented, and perhaps merit recognition as a separate

type of microhair, the ‘pentameroid’ type.

Abaxial macrohairs

Abaxial macrohairs were only observed on the abaxial

surface of some specimens of Pentameris distichophylla,

P. hirtiglumis and P. glacialis. In all these taxa, the hairs

were long, usually produced from the intercostal regions,

had a distinctly swollen base and were surrounded by four

or more cells, termed ‘modified cells’ by Ellis (1986).

The taxonomic value of these structures is limited, as it

30

Bothalia 23,1 (1993)

appears that they are not universally present throughout

all specimens examined of all taxa. Inconsistent sampling

(variation in the region of the leaf from which the samples

were taken, or the age of the leaf blade) may explain the

lack of macrohairs in some samples.

Adaxial macrohairs

Two different hair-like structures were found on the

adaxial leaf surface in some of the taxa of Pentameris,

sometimes both occurring on the same leaf. Conventional

macrohairs, those which have a distinct basal structure

comprised of modified cells, were frequently observed on

the adaxial epidermis in Pentameris thuarii, P glacialis

and P. distichophylla , but were only occasional in P.

swartbergensis. Figure IF illustrates these hairs on the

adaxial leaf blade surface in P thuarii.

The second type of macrohair, termed here a ‘filament’,

is a long, unicellular hair without any obvious basal

differentiation or associated cells. Although no measure-

ments were taken, the filament type of macrohair appeared

to be shorter than the conventional type. Filaments were

frequently observed in the bottom of the adaxial furrows

in specimens of P macrocalycina and occasionally in P.

distichophylla and P. swartbergensis. In P. oreophila

filaments were only found in the furrow nearest the edge

of the lamina. One of these filaments observed in P.

distichophylla is shown in Figure 2A.

Adaxial macrohairs are of slightly greater taxonomic

significance than abaxial macrohairs. The presence and

type of macrohair are useful characters in Pentameris,

although some variability within the species exists. It

appears that the species with open or folded leaves (P.

thuarii, P. distichophylla and P. swartbergensis) possess

conventional macrohairs, whereas those taxa with perma-

nently rolled or folded leaf blades have the second type

of macrohair (P. macrocalycina, P. oreophila, P. disticho-

phylla and P. swartbergensis ). P. swartbergensis and P.

distichophylla thus have both types of macrohair. It is

possible that the filament macrohairs are conventional

macrohairs that have lost the modified cells around the

hair base. The loss of these modified cells may be related

to the evolution of the permanently rolled leaf blade, where

the adaxial surface is not as exposed to the environment.

Prickles

The adaxial surfaces of all the specimens examined had

prickles of one form or another. These different prickle

types (Table 1) are invariant within the taxa of the genus.

FIGURE 2.— Leaf blade SEM of Pentameris. A, P. distichophylla (Lehm.) Nees, adaxial surface; B, P. oreophila N.P. Barker, adaxial surface,

note extreme length of prickles and presence of collar at base of each prickle; C, P. longiglumis (Nees) Stapf subsp. gymnocolea N.R

Barker, adaxial surface, note extreme density and inclination of prickles; D, P. uniflora N.P. Barker, adaxial surface showing prickles

and small microhairs. C, collar; P, prickles; Ma, macrohairs; Mi, microhairs; S, stomata. Scale bars: A, 48 /cm; B, 50 ^m; C, 87 fun;

D, 120 ^m.

Bothalia 23,1 (1993)

31

Characters such as prickle density, distribution, orienta-

tion, length and structural differences are all taxonom-

ically useful for distinguishing taxa within Pentameris

(Figure 2A— D).

The prickles of P. oreophila are long and erect, with

the tips sometimes bent or recurved. In addition to

their unusual length, the prickles possess a basal collar-

like structure (Figure 2B), a character shared with P.

hirtiglumis. The prickles of the latter species are, however,

much shorter, and are almost appressed to the surface of

the leaf, a character shared with P. longiglumis (Figure

2C).

Of the remaining species, P. macro calycina, P. thuarii,

P. glacialis and P. swartbergensis have short, erect

prickles, the tips of which may be bent or curved; are

swollen basally, and the sides are convexly shaped; are

densely distributed, particularly so in P macrocalycina.

The prickles of P. uniflora are also quite densely

distributed and almost erect, as indicated in Figure 2D.

In contrast to the above, P. distichophylla has erect

prickles that are almost straight-sided (as opposed to the

slightly inflated convex sides of the prickles of the other

taxa); they appear to have knobbed tips (Figure 2A) and

are sparsely distributed over the adaxial surface, usually

only on the edges of the ribs. Both the short, inflated

prickle type and the erect, knobbed prickle type have also

been observed in species of Pentaschistis, for example

P. colorata (Ellis & Linder 1992). The long, collared

prickles observed in P. oreophila are unique among the

African Arundineae.

CYTOLOGY

Preliminary cytological studies have shown that

Pentameris distichophylla has a chromosome count of

2n=36, whereas P. thuarii has a count of 2n=12. The

latter figure has been corroborated by H. du Plessis

(pers. comm.). Both these counts were obtained from

meiotic material. P. distichophylla is therefore hexaploid.

Such polyploidy is not unusual in the southern African

Arundineae (see for example counts for Merxmuellera,

Pentaschistis and other genera given by Du Plessis & Spies

1988; Spies & Du Plessis 1988; Spies et al. 1990).

These counts further support the contention that the base

chromosome number for the Arundineae appears to be

x=6 (Davidse 1988), and not x=12 as proposed by Clayton

& Renvoize (1986). Despite the apparent similarities

between Pentameris and Pentaschistis, the different base

chromosome number of the latter genus (x=7 in many of

the known instances) does not suggest a close relation-

ship between these two taxa. It is, however, possible that

Pentameris evolved by means of aneuploidy from a

pentaschistoid ancestor.

PHYLOGENY

The data presented in Table 1 were converted into a data

set suitable for cladistic analysis. The anatomical

characters were augmented by a few morphological

characters, resulting in a data set of 26 characters

(presented in Tables 2 & 3). Four of these characters are

TABLE 2. — Characters used in the cladistic analysis. 1’ = primary

vascular bundles, 3’ = tertiary vascular bundles

Leaf blade characters (t/s)

1. Leaf blade shape in t/s: 0 = flat to curled, 1 = rolled,

2 = permanently rolled

2. 1’ rib shape: 0 = squared, 1 = round

3. 3’ rib shape: 0 = squared, 1 = round, 2 = conical

4. Relative rib depth: 0 = 1’ = 3’, 1 = T <3’

5. Furrow depth: 0 = shallow (<'/2 lamina thickness) 1 = deep

6. Furrow width: 0 = narrow, 1 = wide

7. 1’ Abaxial sclerenchyma girder: 0 = discrete, 1 = continuous

with hypodermis

8. Sclerenchyma cap on leaf margin: 0 = discrete, 1 =

continuous with hypodermis

9. Leaf blade thickness: 0 = uniform thickness, 1 = swollen

near margin

10. Abaxial epidermal cells: 0 = small (< = 25 /im wide),

1 = inflated

11. Silica body shape: 0 = round, 1 = dumbbell-shaped

Adaxial epidermal characters

12. Conventional macrohair type: 0 = absent, 1 = present

13. Filament macrohair type: 0 = absent, 1 = present

14. Prickle type: 0 = inflated basally, 1 = markedly elongated,

2 = apically knobbed

15. Prickle orientation: 0 = erect, 1 = inclined/appressed

16. Prickle base: 0 = unomamented, 1 = collar present

17. Prickle density: 0 = sparse, 1 = dense

18. Microhair type: 0 = basal cell length » apical cell length

1 = basal cell length = apical cell length

19. Microhair length: 0 = short ( < 60 jim), 1 = long

20. Papillae: 0 = absent, 1 = present

Morphological characters

21. Growth form: 0 = unbranched, 1 = branched

22. Glands: 0 = absent, 1 = present

23. Lodicule margin: 0 = entire, 1 = ciliolate

24. Achene surface: 0 = rugose, 1 = colliculate, 2 = reticulate

25. Fruit type: 0 = caryopsis, 1 = achene

26. Ovary appendages: 0 = absent, 1 = present

multistate, and are treated as undirectional in the analysis.

The ‘ie’ option of the cladistic package HENNIG86 version

1.5 was used to analyse the data. Using this method, the

complete set of most parsimonious trees is found (Platnick

1989).

Choice of outgroup

Pentaschistis is the obvious outgroup, although the

recognition of a particular species or group of species

within this genus as the closest to the study group is not

possible on an a priori basis. Six species of Pentaschistis

were therefore chosen as possible outgroup taxa. These

taxa were chosen on the basis of inferences made by Ellis

in his anatomical papers (1985c, 1986), and after discus-

sions with Dr H. P. Linder. The taxa chosen were P. aspera

(Thunb.) Stapf, P. colorata (Steud.) Stapf, P. curvifolia

(Schrad.) Stapf, P. eriostoma (Nees) Stapf, P. glandulosa

(Schrad.) Linder and P. tortuosa (Trin.) Stapf.

Two of these taxa, the glandular P. glandulosa and P.

aspera, were included because they were considered to

be most dissimilar to the study group. Following the

outgroup substitution method of Donoghue & Cantino

(1984), these two least related glandular taxa were used

as the initial outgroup to determine the relationships

between the remaining taxa. Using this method, the single

32

Bothalia 23,1 (1993)

TABLE 3. —Data used in cladistic analyses. Characters marked by asterisks above the data set were removed in the data

set where Pentaschistis curvifolia was used as the outgroup (data set delimited below the dashed line). Characters

appear in the order presented above. The question mark (?) indicates absent data

or few non-study group taxa found to be most closely

related to the study group were then used to perform the

final cladistic analysis on the data set from which the other

outgroup taxa had been removed.

P. swartbergensis share a flat leaf blade with wide furrows

between the adaxial ribs (char. 6). P. glacialis and P.

thuarii share a character that has arisen in parallel else-

where: entire lodicule margins (char. 23).

When the two distantly related taxa are used as the out-

group, one shortest length tree (1. = 60, c.i. = 50, r.i.

= 67) is generated. In this tree (not presented), P. cur-

vifolia is the taxon most closely related to the study group,

and was used as the outgroup (the other Pentaschistis spe-

cies were removed from the data set); the data were re-

analysed after the autapomorphies and invariant charac-

ters had been removed. Once again, HENNIG86 found

one shortest tree, presented in Figure 3 (1. = 34, c.i. =

64, r.i. =70). The phylogeny of the study group in this

tree is identical to that obtained in the initial tree (not

presented) where the two glandular taxa are used as an

outgroup.

Character distribution

In discussing the distribution of the characters on the

tree, the characters from the complete database are used,

although in the smaller data set (where Pentaschistis

curvifolia is the outgroup) some are excluded because they

are constant or autapomorphic. The distribution of the

characters is shown on this tree (Figure 3).

The genus Pentameris is a monophyletic clade supported

by the ovule and fruit characters; the apical appendages

and the achene fruit type (char. 25 and 26, represented

by the star in Figure 3).

Within Pentameris, two smaller clades are recognised.

The first clade comprising P distichophylla, P. thuarii,

P. glacialis and P swartbergensis is supported by the

presence of conventional macrohairs on the adaxial surface

(char. 12). Within this group, P. distichophylla is basal

to the other three taxa and is characterised by the

autapomorhies of knobbed prickles (char. 14) , the rounded

ribs of the tertiary vascular bundles (char. 3) and the rugose

pericarp surface (char. 24). P. thuarii, P glacialis and

The other clade within the genus is supported by the

presence of dense prickles (char. 17), as well as other

characters which undergo reversals in some of the lower

P. glacialis

P. thuarii

P. swartbergensis

P. distichophylla

P. uniflora

P. longiglumis

P. hirtiglumis

P. macrocalycina

P. oreophila

FIGURE 3. — Shortest cladogram produced by HENNIG86 using the

‘ie’ option. The outgroup used was Pentaschistis curvifolia

(Schrad.) Stapf. Length = 34, c.i. = 64, r.i. = 70. Numbers

represent characters (given in Tables 2 & 3; ‘a’ = state 0, ‘b’

= state 1 and ‘c’ = state 2). Solid bars represent apomorphies,

parallel lines represent parallelisms and crosses represent character

state reversals. Star at base of Pentameris clade represents

apomorphic fruit characters: achene fruit type and presence of

apical hairs on ovary.

Bothalia 23,1 (1993)

33

clades (char. 10 and char. 13: abaxial epidermal cell width

and presence of filament macrohairs respectively). P.

unijlora is basal to this clade, characterised by the presence

of one floret (data not included in data set) and short micro-

hairs (char. 19). The remaining four taxa are united

by the presence of a continuous hypodermal band of

sclerenchyma which connects to the abaxial girders and

cap on the leaf margin (char. 7 and 8), and primary ribs

which are deeper than the tertiary ribs (char. 4). P.

longiglumis and P hirtiglumis both have enlarged or

swollen leaf blade margins (char. 9), as does the outgroup.

P. macrocalycina and P. oreophila share conical tertiary

ribs (char. 3) and permanently rolled leaf blades (char.

1). The latter species is distinguished by the autapomorphy

of long prickles (char. 14).

The low consistency index obtained from the analysis

of the complete data set (c.i.=50) indicates the homoplasy

present within the data set. This is further demonstrated

by the increase in the value of the consistency index with

the removal of all but the closest outgroup taxa (c.i. =64).

Characters which are synapomorphies in the reduced data

set are often homoplasic in the complete data set. For

example, the unequal primary and tertiary vascular bundle

thicknesses (char. 4) is a synapomorphic character for the

P. longiglumis, P hirtiglumis, P. macrocalycina and P.

oreophila clade in the analysis based on the reduced data

set. However, in the large data set, this character is also

present in Pentaschistis aspera, P. glandulosa, P. colorata

and P. tortuosa. Similar situations are found in other

characters. Such homoplasy has probably been one of the

root causes bedevilling the efforts of taxonomists, both past

and present, to find an equitable classification for this

group.

Taxonomic implications

The monophyletic status of Pentameris is supported by

the fruit and ovary characters. However, as Pentameris is

undoubtedly closely related to Pentaschistis (at least the

eglandular taxa), the possibility that the latter genus is

paraphyletic ought not to be ruled out. The observation

that Pentameris was placed within, and terminal to, the

Pentaschistis clade in the analysis of the complete data set

provides some evidence as to this possibility. If this is the

case, the name Pentameris would be retained for the

monophyletic assemblage containing the taxa with the

unique fruit characters, while other genera may have to

be erected to accommodate monophyletic groups within

the Pentaschistis assemblage. Only a detailed phylogenetic

study of both Pentaschistis and Pentameris can provide

sufficient data to properly test any hypothesis of paraphyly.

CONSERVATION STATUS

Of the previously known taxa of Pentameris , only P.

obtusifolia is listed in the Red Data Book as a rare,

threatened or extinct species, where it is listed under the

synonym ‘P. squarrosa ’ (Hall & Veldhuis 1985). The

conservation status for this taxon is given as ‘uncertain’.

However, as discussed above, the nomenclature of this

taxon has been found to be confused, and P. obtusifolia

is not included in the genus Pentameris.

Certain species of Pentameris nonetheless require

consideration and recognition as taxa requiring conserva-

tion. Most of the new taxa described below are known from

only a few localities, and should perhaps be regarded as

‘rare’. However, as many of these taxa are found in

inaccessible and thus rarely collected areas, the true

distribution is not fully known. In addition to the problem

of varying collecting intensity, the effects of fire on the

germination, growth and re-establishment of fynbos grass

species are poorly known. Linder & Ellis (1990a) discuss

the various strategies that fynbos grasses have evolved to

escape or adapt to fire. These authors note that grasses

appear in abundance in the first few years after fire, but

are then almost inevitably outcompeted by members of the

Restionaceae and woody fynbos elements. The collection

of fynbos grasses is therefore best carried out in areas that

have been burnt within the previous few years. Such areas

are not always attractive to collectors intent on obtaining

other woody or non-graminoid taxa. Mature, woody fyn-

bos may therefore hide many new or apparently rare grass

species which are present as dormant seeds or under-

ground vegetative structures. Actual species distributions

may therefore be wider than presently known.

In assessing conservation status of the taxa of Penta-

meris, the categories proposed by Rabinowitz (1981) and

modified by Karron (1987) will be used. These categories

record the geographic distribution, demographic structure

and habitat requirements in a two-state form: restricted

or widespread, sparse or abundant, wide or narrow respec-

tively. The demographic component can thus reflect, to

a certain extent, the biological aspects of rarity of the taxa.

As the category of geographic distribution is scale

dependent, it is used here at the level of magisterial

districts or mountain ranges, whichever is more geographi-

cally homogeneous. The allocations of the species of

Pentameris to these categories is presented in Table 4.

Detailed descriptions of distributions and habitats are

provided under individual species descriptions.

TAXONOMY

Taxonomic descriptions of all the taxa are presented.

The dimensions given below for the lemma body, lemma

TABLE 4.— The modified Rabinowitz conservation categories allocated

to each taxon of Pentameris. The allocation of these codes is based

on field observations and data obtained from labels on herbarium

specimens

A = abundant; N = narrow; R = restricted; S = sparse; W = wide;

WS = widespread.

34

Bothalia 23,1 (1993)

FIGURE 4 .—Pentameris uniflora N.P. Barker. A, habit with decumbent, branching culm basally covered in dead, appressed leaf sheaths;

B, ligule, which is a row of hairs, and surrounding leaf parts; C, panicle subcontracted to lax, with few spikelets. D— G, parts of spikelet

and florets: D, glumes; E, lemma (in its normal conformation) showing lemma bristles, lobes (which are adnate to the bristle for most

of their length) and geniculate awn; F, palea; G, ovary with apical hairs and stigmas. A, x 0.5; B, D-F, X 4.7; C, X 1.6; G, x 11.8.

Bothalia 23,1 (1993)

35

lobes and lateral bristles are measured as follows: the

lemma body is measured from the base to the lowermost

point of insertion of the central awn, whereas the lobes

and bristles are measured from this same insertion point

to the apex of the lobe or bristle. The full length of the

lemma (either from base to lobe apex or base to bristle

tip) can be obtained by adding the lemma body length to

the length of the lobe or bristle. The term ‘culm’ is used

in the descriptions below to describe all aerial parts

(branched stems and reproductive culms) despite the

anatomical discrepancies discussed in the introduction.

Pentameris Beauv. , Essai d’une Nouvelle Agrosto-

graphie: 92, t.18, fig.8 (1812); Kunth: 107 (1829); Kunth:

315 (1833); Nees: 336 (1841); Stapf: 512 (1897); Chippin-

dall: 251 (1955); Clayton & Renvoize: 174 (1986); Gibbs

Russell et al. : 251 (1990). Type species: Pentameris thuarii

Beauv.

Perennial; caespitose, cushion-like or decumbent, some-

times branched. Culms 250—2 000 mm in length; woody

and persistent from a woody base; leaf sheaths appressed

to culm or free, persistent; sheath mouth sometimes auric-

ulate; ligule a fringe of hairs; leaf blades linear, 50—550

mm long, rigid or filiform, open and flat, rolled or

acicular; inflorescence paniculate, lanceolate and some-

what contracted to globose and lax; spikelets solitary,

13-25 mm long, laterally compressed, 1-2-flowered,

highly reduced third floret rarely present; glumes two,

more or less equal, 1-nerved, membranous to chartaceous,

minutely scabrid, sometimes hirsute; lemma body uni-

formly pubescent, 7— 9(— ll)-nerved, 2.2— 6.0 mm long,

pubescent, the hairs arising from between the nerves,

nerves anastomosing into the awn base and a 2.5-12.0 mm

long lateral bristle; lemma lobes 0.4-4.0 mm long, acute

to acuminate, sometimes lacerated, partly to fully adnate

to the bristles; central awn geniculate, scabrid, contorted

basally, 2—11 mm long from base to knee, 6—21 mm long

from knee to tip; palea longer than lemma body, bifid at

apex, pubescent between the keels; lodicules two, glabrous

or ciliolate, sometimes with arm-like extensions; stamens

three; ovary apically hairy, these hairs retained until matu-

rity; fruit an achene with free pericarp, fusiform, subglo-

bose to globose or cuneate, surface colliculate or rugose,

hilum up to two thirds the length of the fruit, embryo

small.

A genus of nine species, endemic to the southwestern

region of the Cape Province, South Africa.

Key to species

La Spikelets with one floret 1. P. uniflora

lb Spikelets with two florets, occasionally with a rudimentary third floret:

2a Leaves with purple to dark brown auricles at base of blade; lemma lobes truncate, apically dentate .. 2. P. thuarii

2b Leaves without auricles, or if present then not coloured as above; lemma lobes acute to acuminate:

3a Panicle lax at anthesis, globose, 170-300 mm long; basal leaf sheaths 120 mm or longer, clustered at,

and free from culm base; culm, including inflorescence, usually taller than 1.2 m:

4a Leaf sheaths densely covered in short hairs; glumes 15—18 mm long; lemma body 3.0— 3.5 mm long; known

only from Table Mountain 3a. P. longiglumis subsp. longiglumis

4b Leaf sheaths glabrous; glumes 21—24 mm long; lemma body 5.0— 5.5 mm long; known only from the Kogel-

berg 3b. P. longiglumis subsp. gymnocolea

3b Panicle subcontracted to contracted at anthesis, lanceolate, up to 150 mm long; basal leaf sheaths seldom

longer than 120 mm, partially free or appressed to culm; culm, including inflorescence, seldom taller than

1.2 m;

5a Glumes 14—24 mm long; leaf blades acicular or permanently rolled, rigid, sometimes pungent:

6a One or both glumes pubescent 4. P. hirtiglumis

6b Glumes never pubescent:

7a Leaf blades acicular, usually straight but sometimes curling toward apices, occasionally pungent; leaf

sheaths glabrous, but sheath mouth may be bearded; sheaths closely appressed to culm . 5. P macrocalycina

7b Leaf blades permanently rolled, usually falcate, strongly pungent; leaf sheaths not appressed to culm,

always pubescent, if only at margins 6. P. oreophila

5b Glumes usually 14 mm or shorter; leaf blades usually soft, rolled (not permanently), folded or flat in cross

section, never pungent:

8a Lateral bristle of lemma 5.5-10.0 mm long; leaf sheaths pubescent to densely woolly, especially near sheath

mouth 7. P. distichophylla

8b Lateral bristle of lemma 2. 5-5.5 mm long; leaf sheaths glabrous or pubescent at margin only:

9a Leaves 200 mm long or longer, up to 5 mm wide; panicle with 25 or more spikelets 8. P. swartbergensis

9b Leaves usually shorter than 100 mm, thin and rolled, narrower than 2 mm; panicle with fewer than 15

spikelets 9. P. glacialis

1. Pentameris uniflora N.P. Barker, sp. nov.

A aliis omnibus speciebus Pentameridis flosculo singu-

lari differt.

Pentameris sp. 2. in Gibbs Russell et al.: 253 (1990).

TYPE. — Cape, Riversdale, Sleeping Beauty Peak.

Along edges of overgrown ledges or on steep south slopes.

Dense. 4200ft (1 275 m), 29 Oct. 1967, Esterhuysen 31771

(PRE, holo. ! ; BOL, iso.!).

Plants soft, decumbent; culms thin and flexuous, up to

650 mm long; leaf sheaths glabrous or only pubescent

along margins, appressed to culm; leaf blades soft,

filiform, short, up to 125 mm long, open or folded to some-

what rolled; panicle lanceolate, delicate, 40—60 x 10—20

mm, lax; spikelets 5—20, one-flowered; glumes 11—12 x

0.9— 1.0 mm; lemma body 5.5— 6.0 mm long; lemma lobes

acuminate, 1. 5-3.0 mm long, adnate to 2. 5-4. 5 mm long

lateral bristle for most of their length; awn geniculate,

2.0— 3.5 mm from base to knee, 6.5— 7.0 mm from

knee to tip; palea 5.5— 6.0 mm long; lodicules cuneate,

glabrous; anthers 4 mm long; fruit not known; flowering

time September to December. Figures 2D; 4.

36

Bothalia 23,1 (1993)

FIGURE 5. — Distribution of P. glacialis N.R Barker, ■; P. hirtiglumis

N.P. Barker, O; P longiglumis (Nees) Stapf subsp, longiglumis

N.P. Barker, ▼ ; P. longiglumis (Nees) Stapf subsp. gymnocolea

N.P. Barker, V ; P oreophila N.P. Barker, #; P. swartbergensis

N.P. Barker, *; P uniflora N.P. Barker, ♦; neighbouring locality

of P. hirtiglumis and P. oreophila, C).

This species is the only taxon in the genus with one-

flowered spikelets. Mature fruit have not been seen, but

stylar hairs on the apex of the ovary confirm the position

of this species in the genus Pentameris (Barker 1990).

P uniflora is known from only three localities which

are quite widely separated (Figure 5). Populations at these

localities appear to be quite abundant, but the habitat of

this species (damp, rocky, southern aspects of the Cape

fold mountains), habit (decumbent, hidden under taller

plants), and small, somewhat inconspicuous inflorescences

may account for the paucity of herbarium specimens.

Vouchers: Bond 1581 (SAAS); Ellis 2546 (PRE); Esterhuysen 25025,

32718a (BOL); Esterhuysen 31771 (BOL, PRE).

2. Pfentameris thuarii Beauv., Essai d’une Nouvelle

Agrostographie: 92, t. 18, fig. 8 (1812); Roem. & Schult.:

693 (1817); Kunth: 107 (1829) as ‘thouarsii’; Kunth: 315

(1833) as ‘thouarsii’; Kunth: 270 (1835) as ‘thouarsii’; Stapf:

513 (1897); N.P. Barker in Gibbs Russell etal.: 253. (1990).

Type: Du Petit-Thouars s.n. (P, lecto.!). Lectotype here

designated.

Danthonia thuarii Desv. : 99 (1831); Nees: 337 (1841) as ‘thouarsii’;

Steudel: 243 (1855) as ‘thouarsii’; Durand & Schinz: 854 (1895) as

‘thouarsii’; Durand & Schinz: 854 (1895).

P thuarii Beauv. var. burchellii Stapf: 513 (1897). Type: Riversdale

Div., lower part of the Lange Bergen, near Kampsche Berg, Burchell

6964 (K, holo.!; GRA, iso.!; PRE (fragment), iso.!).

Plants caespitose or decumbent; culms 0.46—1.63 m

long; leaf sheaths pubescent, appressed to culm, with

purple auricle at mouth, persistent; leaf blades up to 500

mm long, folded or flat (rarely rolled); panicle globose,

lax, 70—220 x 40—170 mm; spikelets 16—90, two-

flowered; glumes 15.5—21.5 x 1.8— 3.0 mm; lemma body

2.2— 3.3 mm long; lemma lobes 0.4— 0.7 mm long, acute,

apically lacerate-dentate, almost free from a 4.0— 5.5 mm

long lateral bristle; awn geniculate, 4.5 —7.0 mm from base

to knee, 9.5 — 12.5 mm from knee to tip; palea 2.8— 3.5

mm long; lodicules cuneate, glabrous; anthers 3—4 mm

long; fruit an achene, globose, 2.5 x 1.7— 2.0 mm, surface

colliculately sculptured; flowering time September to

December. Figure IB, D, F.

The spelling of the epithet thuarii, as used in the proto-

logue, differs so obviously from that of the name of the

collector, Du Petit-Thouars, that it is not considered a

typographic error. The epithet thouarsii, used in a number

of subsequent accounts, is therefore not adopted. It is

plausible that Palisot de Beauvois wished to maintain the

phonetic pronunciation of the French name ‘Thouars’, and

thus spelt it ‘thuarii’ in latinised form.

No locality or specimen number of the type specimen

was cited by Palisot de Beauvois. However, two specimens

of P. thuarii were obtained from the Paris herbarium (P).

One of these specimens is annotated with ‘Herb. Du Petit

Thouars’, which is written on the bottom of the label. The

identity of the handwriting is not known. That one (or

both) of these specimens are type material is further

supported by the observation that the illustration accom-

panying the original description includes a representation

of a mature fruit, a description of which also appears in

the text (Palisot de Beauvois 1812). The type specimen

must therefore have borne mature fruit. This deduction

is compatible with the historical record which indicates

that Du Petit Thouars visited the Cape in Febuary 1793.

February is the time of year when many fynbos grasses,

including P thuarii, are in fruit (pers. obs.). As the

annotated specimen from Paris is in full fruit, this

specimen is regarded as type material. However, this

evidence is considered to be insufficient to warrant

holotype status for this specimen, which is therefore

selected as a lectotype.

Stapf (1897) distinguished two varieties of P thuarii : var.

thuarii, was described as being only 1—2 ft. (300—600

mm) tall with a woody, sometimes branched, suffrutes-

cent base, with leaves only 1 lin. (approximately 2 mm)

wide and with a lax, open or contracted panicle; var.

burchellii Stapf was considered to be taller, simple basally

Bothalia 23,1 (1993)

37

and branching after 1 ft. (300 mm), with leaves 2 lin.

(approximately 4-5 mm) wide and with an effuse panicle.

Stapf added that this latter variety has the appearance

of a robust annual. Chippindall (1955) considers the

characters on which these distinctions are made to be too

variable, and rejects the two varieties, a conclusion

supported by Barker (1990).

This species grows abundantly in moist environments,

and is usually found alongside streams, seeps and drainage

lines. Geographically widespread, it occurs from Stellen-

bosch in the west to Montagu Pass in the east (Figure 6).

Vouchers: Ellis 2221 (PRE); Compton 7494 (BOL, NBG); Levyns 693

(BOL, STE); Parker 4465 (BOL, NBG, SAM); Schlechter 9282 (BOL,

GRA, PRE).

3. Pentameris longiglumis (Nees) Stapf

The following description is based on all the specimens

seen, and the variation reported includes that for both

known subspecies, described below.

Plants densely tufted from a woody base; culms erect,

up to 1.4 m tall, unbranched or branched close to woody

base; leaf sheaths loose and free from culm, wide,

persistent, clustered at base of plant, shortly pubescent

or glabrous, straw-coloured or purple, sheath mouth some-

times with green auricles; leaf blades rigid, rolled, up to

550 mm long; panicle lax, globose, up to 300 x 140 mm;

spikelets 30—100 or more, two-flowered; glumes 15.0—

24.0 x 1.1— 2.4 mm; lemma body 3.0— 5.5 mm long;

lemma lobes 2. 4 -4.0 mm long, long -acuminate, almost

completely adnate to a 3.5—11.0 mm long lateral bristle;

awn geniculate, 3.5—10.0 mm from base to knee, 7.5—21.0

mm from knee to tip; palea 3.0— 9.5 mm long; lodicules

cuneate, glabrous; anthers 3.6-5. 5 mm long; fruit broadly

fusiform, 3.5 x 1.3 mm, surface colliculately sculptured;

flowering time September to December. Figures 8 & 9.

Two subspecies are recognised on the basis of a number

of floral and vegetative characters, as listed in Table 5.

The significance of these differences is demonstrated by

the results of a principal components analysis (PCA)

TABLE 5. — A statistical summary of the variation in characters

differentiating the two subspecies of P. longiglumis (Nees) Stapf.

Measurements in mm

carried out on data obtained for 11 characters from 10

herbarium specimens. The PCA was done using the

numerical taxonomy package NT-SYS, version 1.4 (Rohlf

1988).

The first three axes resulting from the PCA accounted

for 94.2% of the variation in the data, with the first axis

contributing 75.7 % . The first two axes (Figure 7) account

for 88.3% of the variation in the data. The two groups

comprise specimens from two geographically distinct

localities, Kogelberg and Table Mountain. On the basis

of this evidence, the allocation of the rank of subspecies

to these two groups is considered to be justified.

3a. Pentameris longiglumis (Nees) Stapf subsp.

longiglumis

Danthonia longiglumis Nees in Flora africae australioris I: 306 (1841).

TYPE.— Cape, In summo monte tabulare (Dist. Cap.),

Bergius s.n. (B, holo.!; fragment and photo in PRE, iso.!).

Note: Pentameris longiglumis sensu Stapf: 514 agrees

with neither Nees’s description nor the type (see discus-

sion below).

0.6 -

*\ Tay 7231

A

x o.i

subsp. longiglumis

o\

Mar 3063

O Be 15170

Mar 3078

-0

V

O Bark 994

\

Bark 993

Bou 1649

SZ 11292

Ell 2341 /

v---' subsp. gymnocolea

n — i — i — i — i — i — i — i — i — i — i — i — i — — i — i — i — i — i — i — i — i — i — i — i — i — i — i —

-1.2 -1.0 -0.8 -0.6 -0.4 -0.2 0 0.2 0.4 0.6 0.8 1.0 1.2 1.4

Axis 1

FIGURE 7. —First two axes of the

principal components analysis

on data of eight specimens of

Pentameris longiglumis (Nees)

Stapf. Axis 1 accounts for

75.7 % of variation in data, axis

2, 12.6%. Points represent

specimens, annotated as fol-

lows: Bark = Barker, Be -

Bews, Ell = Ellis, Mar =

Marloth, SZ = Stirton &

Zantovska, Tay = Taylor, VanR

= Van Rensburg. Specimens

from BOL, PRE and STE.

38

Bothalia 23,1 (1993)

FIGURE 8 —Pentameris longiglumis (Nees) Stapf subsp. gymnocolea N.P. Barker. A, habit: erect culms, basally covered in loose, dead leaf

sheaths; note long leaf blades, which may be rolled or open, and large, globose, effuse panicle, x 0.8; B, ligule, which is a row of hairs,

surrounding leaf parts, with hairs on leaf sheath, expanded auricles, and sparsely pubescent adaxial basal leaf surface, x 4.7.

Bothalia 23,1 (1993)

39

FIGURE 9. — Pentameris longiglumis (Nees) Stapf subsp. gymnocolea

N.R Barker. Parts of spikelet and florets: A, long, acuminaie

glumes; B, whole basal floret; C, lemma (opened, flattened and

viewed from inside of floret) indicating venation, lemma bristles,

lemma lobes (adnate to bristles for most of their length) and

geniculate awn; D, palea; E, developing fruit, with apical hairs.

A-D, X 3.7; E. x 11.

Stapf (1897) describes P longiglumis as only 1.5 to 2

feet (450-600 mm) tall. Nees does not give any indica-

tion of plant height, but all specimens of this taxon seen,

including the type (Bergius s.n.), have far exceeded this

height. Furthermore, Stapf describes the sheath as glabrous

or sparsely hairy, whereas the type specimen has a

distinctly pubescent sheath, as described by Nees (1841).

In addition, Stapf describes the leaves as being filiform,

6—9 inches (150—220 mm) long. Nees does not provide

leaf lengths in his description, but leaves on the type

specimen were noted to be much longer than 6—9 inches,

broad, rolled and somewhat rigid.

Furthermore, at least three of the four specimens cited

by Stapf under his description of P. longiglumis do not

match the Bergius specimen cited by Nees. The three

specimens in question, Burchell 542, Burchell 598 and

Milne 246 were obtained from The Herbarium, Royal

Botanic Gardens, Kew (K). The fourth voucher cited by

Stapf, Spielhaus s.n. ,was not traced. Unfortunately these

three specimens are of poor quality. The spikelets (when

present) are devoid of florets, and one specimen, Burchell

542, is entirely sterile. Uncertainty about the identity of

the Milne 246 specimen is further borne out by an anno-

tation to the effect that the leaves resemble those of

Pentaschistis pallescens (Schrad.) Stapf, whereas the

inflorescence resembles that of Pentameris thuarii. The

Burchell 598 specimen may possibly be Pentameris longi-

glumis as it has floral parts of a size comparable to those

observed in Nees’s type specimen . However, the basal parts

are absent, so the leaf and sheath characters cannot be

compared.

It is therefore considered that Stapf’s (1897) description

and cited vouchers do not agree with Nees’s description

and type of P. longiglumis . Stapf s Pentameris longiglumis

is therefore regarded as a misapplied name, and it appears

that Stapf described a species of Pentaschistis.

This subspecies is known from a few specimens

collected from Table Mountain, the distribution shown in

Figure 5. Until recently, this taxon was thought to be

extinct, as the most recent specimen seen among the

holdings of eight South African herbaria is dated 1918.

However, a small population has been located in a recent-

ly burnt area on Table Mountain. This population is

growing in a slightly sloping seepage area. The moist

habitat requirement matches that of the other subspecies,

which is, however, found on steeper slopes.

Vouchers: N. Barker 993, 994 (BOL, PRE); Bews 15170 (BOL);

Marlorh 3063, 3078 (PRE).

3b. Pentameris longiglumis ( Nees ) Stapf subsp.

gymnocolea N.P. Barker, subsp. nov.

Differt a P. longiglumi subsp. longiglumi foliorum vagi-

nis glabris et partibus floralibus grandioribus: glumae

21—24 mm longae, lemma 5.0— 5.5 mm longum; sub-

species tantum monte ‘Kogelberg' nominato nota. Figures

2C; 8 & 9.

TYPE. — Cape, summit ridge of Platberg, Kogelberg,

S side, edge of gully; culms 3—4 ft. (1.0— 1.3 m), erect,

coarse, loose tufts, 2650 ft (900 m), 27 Nov. 1967, Taylor

7231 (PRE, holo.!; STE, iso.!).

As shown in Table 5, this subspecies differs from P.

longiglumis subsp. longiglumis in a number of features.

The subspecific epithet chosen for this species describes

one of these distinguishing characters: the glabrous leaf

sheaths.

This subspecies is known only from the Kogelberg

Forest Reserve (Figure 5). It inhabits south-facing moun-

tain slopes in seepage areas where it forms large tussocks.

Vouchers: Boucher 1649 (PRE); Ellis 2341 (PRE); Esterhuysen 13326

(BOL. PRE); Stirton & Zantovska 11293 (STE); Van Rensburg 518 (STE).

4. Pentameris hirtiglumis N.P. Barker, sp. nov.

P. oreophilae similis sed glumis hirsutis facile distincta;

lobis lemmae acutis, usque ad dimidium longitudinis ad

setam adnatis.

40

Bothalia 23,1 (1993)

FIGURE 10. —Pentameris hirtiglumis N.P. Barker, type specimen Kerfoot 6092. A, habit: decumbent, branching culm basally covered in dead

leaf sheaths; note that flowering culm is not normally curved back upon itself, but appeared so on specimen. B— G, parts of spikelet and

florets: B, hirsute glumes; C, whole basal floret; D, lemma (opened, flattened and viewed from inside of floret) showing venation, lemma

bristles, lemma lobes (adnate to bristle for half or less of their length) and geniculate awn; E, palea; F, anthers, with no filaments; G,

ovary with apical hairs and stigmas. A, x 0.5; B— F, x 4.7; G, x 14.

Bothalia 23,1 (1993)

41

TYPE. — Cape, Bosboukloof, Jonkershoek, locally very

common, attractive, 980 ft (600 m), Oct. 1967, Kerfoot

6092 (PRE, holo.!).

Plants cushion-like or densely bushy, branched; culms

somewhat decumbent with numerous nodes, 200-750 mm

long; leaf sheaths pubescent, at least along margins (rarely

glabrous), persistent, not closely appressed to culm, loose

when dead; leaf blades up to 200 mm long, permanently

rolled, falcate; panicle lanceolate, somewhat lax, 75—110

x 20—35 mm; spikelets 15-50, two-flowered; glumes

hirsute, 14.5—21.5 x 1.6— 2.1 mm; lemma body 2. 8— 3.7

mm long; lemma lobes 1—2 mm long, acute, adnate to

a 3.0-6.5 mm long lateral bristle for half or less of their

length; awn geniculate, 4—6 mm from base to knee,

7.5-11.0 mm from knee to tip; palea 4.0-4. 5 mm long;

lodicules cuneate, glabrous; anthers 3.5— 5.0 mm long;

fruit broadly fusiform, 2.4 x 1.2 mm, surface collicu-

lately sculptured (Figure 1A); flowering time September

to October/November.

This species is similar in appearance to P. oreophila ,

but can be readily separated from that species by the

presence of the hirsute glumes (Figure 10B) . Additional

differences between these two taxa are the acute lemma

lobes (Figure 10D) and softer, non-pungent leaf apices

in P. hirtiglumis. However, as is apparent from the

cladogram discussed above (Figure 3), this species is

sister to P longiglumis on the basis of several anatomical

features including swollen leaf margins, inflated abaxial

epidermal cells and dense, strongly inclined adaxial

prickles.

Like P. oreophila, this taxon has a geographically

restricted distribution, and is known only from high

montane regions of the Hottentots Holland Mts (Figure

5). Where found, this species is abundant and locally

dominant. However, it appears to have narrow habitat

requirements, and is restricted to shale bands.

Vouchers: N. Barker 90 (PRE); Ellis 4680 (PRE); Kruger & Haynes

753 (JF, STE).

5. Pentameris macrocalycina (Steud.) Schweick. in

Feddes Repertorium 43: 91 (1938); N.R Barker in Gibbs

Russell et al.: 252. (1990). Type: Cape, in summitate

Montis Tabularis Ecklon 932 (OXF, lecto.!, PRE isolecto.!

(fragments), S, isolecto.!). Lectotype here designated.

Avena macrocalycina Steud.: 482 (1829).

Danthonia speciosa Lehm. ex Nees: 307 (1841). Steud.: 241. (1855);

Durand & Schinz: 854 (1895). Type: In apicis rupestribus montis ad

Genadenthal solo sabuloso-humoso alt. 2000—3000 ft, (Stellenbosch),

et in Dutoitskloof alt. 2500-3000 ft, Drege s.n. (PRE fragments, iso!.).

Pentameris speciosa (Lehm. ex Nees) Stapf: 515 (1897).

Plants caespitose, branched; culms 0.43—1.10 m long;

leaf sheaths sometimes bearded at mouth, persistent,

closely appressed to culm; leaf blades 110—290 mm long,

acicular; panicle lanceolate, somewhat lax, 60—120 x

10-60 mm; spikelets 10-50, two-flowered; glumes 16-24

x 2-3 mm; lemma body 3.5-5.5 mm long; lemma lobes

1.3— 4.0 mm long, acute-acuminate, adnate to a 7.5—12.0

mm long lateral bristle for most of their length; awn

geniculate, 5.5— 10.0 mm from base to knee, 9— 16 mm from

knee to tip; palea 6.5—11.0 mm long; lodicules cuneate.

glabrous; anthers 4.2— 6.0 mm long; fruit broadly fusi-

form, 3.6 x 1.2 mm, surface colliculately sculptured;

flowering time September to December.

Schweickerdt (1938) pointed out that both Stapf (1897)

and Durand & Schinz (1895) had placed Avena macro-

calycina in synonymy under Danthonia macrantha. In

addition, Nees (1832) mistakenly placed Avena macro-

calycina and its type, Ecklon 932, under Pentameris

macrantha. This confusion arose as a result of the fact

that there were two specimens representing different taxa

distributed under the number Ecklon 932. The type

specimen of Avena macrocalycina, one of the Ecklon 932

specimens, was obtained from the Fielding-Druce

herbarium, Oxford (OXF), and is designated here as the

lectotype. A duplicate of this specimen is housed in the

Swedish Museum (S).

This is a widely distributed species, recorded from

Pakhuis Pass in the north to Cape Point in the south and

eastwards to the Groendal Wilderness Reserve (Figure 11).

Vouchers: Adamson 3980 (JF, PRE); Ellis 2540 (PRE); Esterhuysen

23763, 28012 (BOL, PRE); Taylor 11113 (PRE, STE).

6. Pentameris oreophila N.P. Barker, sp. nov.

Species laminis revolutis permanentibus, brevibus

(<105 mm longis), falcatis, et foliorum vaginis saltern

marginibus pubescentibus distinguitur.

Pentameris sp. 1. in Gibbs Russell et al.: 252. (1990).

P. obtusifolia sensu Ellis (1985d), Barker (1986, 1989). Note that P.

obtusifolia (Hochst.) Schweick. is to be transferred to Pseudopentameris.

TYPE. — Cape, Worcester Div., Jona’s Kop, common

on shale band, or on peaty slopes, after fire, in different

aspects but not on steep southern slopes, forming dense

and, in some places, quite extensive patches, leaf tips very

sharp, 5000 ft (1 500 m), 19 Dec. 1971, Esterhuysen 32681

(PRE, holo.!; BOL, iso.!).

Plants cushion-like or densely bushy, branched basally;

culms erect or somewhat decumbent in older, larger tufts

and then with numerous nodes, up to 530 mm long; leaf

sheaths persistent, pubescent along margins, appressed or

free from culm, especially when dead, often purple when

young; leaf blades up to 105 mm long, permanently rolled,

falcate (less so in young plants), strongly pungent; panicle

lanceolate, somewhat lax, 30—80 x 20—40 mm; spikelets

8—20, two-flowered; glumes 14—20 x 2.1— 3.0 mm;

lemma body 3—4 mm long; lemma lobes 1.6— 3.0(— 4.0)

mm long, acuminate, adnate to 7.0—10.5 mm long lateral

bristle for most of their length; awn geniculate, 6—11 mm

from base to knee, 9.5—13.0 mm from knee to tip; palea

5.5— 8.5 mm long; lodicules cuneate, glabrous; anthers

4.0— 5.5 mm long; fruit subglobose, 3.5— 4.0 x 1.6— 2.0

mm, surface colliculately sculptured; flowering time

September to December. Figures IE & 2B.

The species can be distinguished by its permanently

rolled, short (<105 mm long), falcate leaf blades and the

leaf sheaths which are pubescent at least along the margins.

The specific epithet oreophila, or 'mountain-loving', is

chosen because this species is only found at high altitudes

in the Hottentots Holland, Riviersondereinde and Hex

42

Bothalia 23,1 (1993)

FIGURE 11. — Distribution of P. macrocalycina.

River Mountain ranges (Figure 5). Where found, the

species is abundant, especially in years immediately after

fire. Its high altitude habitat receives snow in the ' /inter

months, and the cushion-like growth form (Figr 2) may

be an adaptation to this.

Vouchers: Ellis 4686 (PRE); Esterhuvsen 19788 (BOL, PRE); Forsvth

191 (JF, PRE); Phillips 2109 (SAM); Wasserfall 562 (BOL, NEG).

7. Pentameris distichophylla (Lehm.) Nees in Lin-

naea 7: 314. (1832); Kunth: 317 (1833). Type: Habitat in

Promontorio Bonae Spei, collector unknown (S, lecto.!,

PRE, fragment).

Damhonia distichophylla Lehm.: 41 (1831); Nees: 305 (1841).

Pentameris dregeana Stapf: 515 (1897); N.P. Barker in Gibbs Russell

et al. : 252 (1990).

Plants caespitose, becoming cushion-like with age,

branched basally; culms 0.4— 1.2 m long; leaf sheaths

pubescent to woolly, especially near mouth, appressed to

culm, persistent; leaf blades 80— 190(— 300) mm long,

pubescent, rolled, folded or flat; panicle lanceolate,

somewhat lax, 50—110 x 15—60 mm; spikelets 12—65,

two-flowered; glumes 12-15 x 1—2 mm, occasionally

sparsely pubescent; lemma body 2.3— 3.5 mm long; lemma

lobes 1.0-2.6 mm long, acute, adnate to a 5.0-10.5 mm

long lateral bristle for most of their length; awn genicu-

late, 4—7 mm from base to knee, 6—12 mm from knee

to tip; palea 4—6 mm long; lodicules cuneate, glabrous

or shortly ciliolate, rarely with arm-like extensions;

anthers 3.0— 4.5 mm long; fruit cuneate, 2.0 x 0.9 mm,

surface rugosely sculptured; flowering time September to

December. Figures 1 & 2.

Stapf (1897) created the epithet dregeana for the partial

concept of the taxon which Nees (1841) called Danthonia

distichophylla. Stapf did not agree with Nees on the

inclusion of Lehmann’s description of Danthonia disticho-

phylla, and in a footnote states that Lehmann’s descrip-

tion probably refers to a Pentaschistis . It thus appears that

Stapf did not see the type of Lehmann’s Danthonia

distichophylla.

Lehmann’s description (1831) states that he has seen the

specimen in a dried state (v.s.= vide siccam). According

to Nordenstam (1980), Lehmann’s herbarium comprised

many specimens collected by Ecklon & Zeyher. The

collections of Lehmann now reside in the Swedish

Museum of Natural History (S), from which the type of

Danthonia distichophylla was obtained. The label of this

specimen had been annotated (possibly by Nordenstam)

to the effect that it is written in Lehmann’s hand. Further

confirmation of this was obtained by matching the script

with that of published examples of Lehmann’s handwriting

(Burdet 1976; Nordenstam 1980). As there are no other

labels on the specimen, the original collector of this

specimen is unknown, but as Lehmann never visited

southern Africa, it is possible that the specimen is an

Ecklon & Zeyher collection.

Despite Stapf s comment, the Lehmann specimen is a

good match to the specimens of P. dregeana cited by Stapf

(1897). The name P. dregeana is therefore illegitimate as

it is antedated by Lehmann’s P. distichophylla. This taxon

is therefore correctly named Pentameris distichophylla

(Lehm.) Nees. As there is no conclusive evidence that the

Lehmann specimen from S is the holotype, it is designated

as the lectotype.

This species is one of the most widely distributed in the

genus, ranging from the northern Cedarberg to Paarl and

eastwards to the Tsitsikamma Mountains (Figure 13).

Vouchers: Esterhuvsen 22352, 27321 (BOL, PRE); Hafstrom & Acocks

45 (PRE); Gillett 3835 (STE); Taylor 11590 (PRE, STE).

Bothalia 23,1 (1993)

43

8. Pentameris swartbergensis N.P. Barker , sp. nov.,

P. distichophyllae similis sed foliorum vaginis glabris

differt.

TYPE. — Cape, Toverkop, Swartberg nr Ladismith,

broad ledges at base of high cliffs on S side of peak,

appears to be locally dominant below cave, 6500 ft. (1 875

m), 17 Dec. 1956, Esterhuysen 26755 (PRE, holo.!; BOL,

iso.!).

Plants caespitose or somewhat decumbent, basally

branched; culms up to 560 mm long; leaf sheaths glabrous,

appressed to culm; leaf blades up to 230 mm long, folded

or rolled, sparsely pubescent near base; panicle lanceo-

late, 80—90 x 20—35 mm, somewhat lax; spikelets

22—56, two-flowered; glumes 11.5—13.0 x 1.5— 1.6 mm;

lemma body 2.8— 3.0 mm long; lemma lobes, acuminate,

1.4— 1.5 mm long, adnate to 2.5— 3.5 mm long lateral

bristle for about half their length; awn geniculate, 3 mm

from base to knee, 5.5 -7.0 mm from knee to tip; palea

4.5— 5.0 mm long; lodicules cuneate, ciliolate at apex;

fruit broadly fusiform, 2.0— 2.4 x 1.0 mm, surface collicu-

lately sculptured; flowering time September to December.

Figure 14.

This species may be confused with P. distichophylla,

but differs in such features as the nature of the colliculate

surface of the caryopsis (which is rugose in P. disticho-

phylla), the leaf sheaths (glabrous in P. swartbergensis,

pubescent to woolly in P. distichophylla) and the lateral

lemma bristles, which are substantially shorter in P.

swartbergensis.

This taxon is known from only two localities in the Klein

Swartberg, and is probably endemic to this mountain range

FIGURE 12. — Pentameris oreophila

N. R Barker. A, habit: erect

culms and prickly, cushion-like

basal leaf growth; note also

pubescent leaf sheaths loosely

appressed to stems, falcate,

permanently rolled leaf blades

with pungent apices. B— G,

parts of spikelet and florets: B,

glumes; C, whole basal floret;

D, lemma (opened, flattened

and viewed from inside of

floret) showing venation, lem-

ma bristles, lemma lobes

(adnate to bristles for most of

their length) and geniculate

awn; E, palea; F, anthers with

no filaments; G, ovary with

apical hairs and stigmas. A, X

O. 3; B-F, x 3.7; G, x 11.

44

Bothalia 23,1 (1993)

(Figure 5). It grows at the foot of cliffs or rock walls in

deep shade.

Vouchers: Esterhuysen 18510, 26750, 26751 (BOL, PRE); Linder 5490

(BOL).

9. Pentameris glacialis N.P. Barker , sp. nov., a P.

distichophylla spiculis paucis (< 15), foliis brevibus filifor-

mibus et habitu decumbenti molli distinguitur.

TYPE. — Cape, Oudtshoom, Waboomsberg, growing in

humic gullies on south slopes, 14 Dec. 1991, N.P Barker

995 (BOL, holo.!; B, CANB, G, GRA, J, K, NBG, NSW,

NU, P, PRE, STE, all iso.!)

Plants decumbent; culms thin, flexuous, up to 550 mm

long, tinged with purple; leaf sheaths glabrous or pubes-

cent along margins, loosely appressed to culm; leaf blades

filiform, short, up to 100 mm long, rolled; panicle

lanceolate, to 70 x 30 mm, lax to somewhat contracted;

spikelets 6— 12(— 15), two-flowered; glumes 13.0—14.5 X

1.3— 1.5 mm; lemma body 2.5— 3.0 mm long; lemma lobes

acuminate, 1.0— 1.9 mm long, adnate to 3.5— 4.5 mm long

lateral bristle for less than half its length; awn geniculate,

3.5— 5.5 mm from base to knee, 7.5— 8.5 mm from knee

to tip; palea 3.5— 4.2 mm long; lodicules cuneate,

glabrous; fruit an achene, 2.5 x 1 mm, surface collicu-

lately sculptured; flowering time October to December.

This species, first collected in 1987 by Ellis, bears

superficial resemblance to P macrocalycina, but is

distinguishable from that species by the slender, geniculate

culms, the rolled but not acicular leaf blades, the panicles

with few spikelets, as well as by the size of the floral

structures, which are substantially smaller in P. glacialis.

The species is generally found to be locally abundant,

growing in narrow gullies, rock ledges and overhangs in

black, humic but sandy soils. Such habitats are thought

to become particularly heavily snowed up in winter. This

habitat appears to preclude P. macrocalycina, which is

found in more stony sites, as well as P. distichophylla,

which appears to prefer northern aspects and rock crevices.

The woolly leaf bases and sheaths of P. distichophylla

separate this species from P. glacialis (Figure 15).

When visited in mid-October 1991, the plants and rocks

at the type locality were covered in a thick layer of ice.

Such conditions may persist for three or more months of

the year in the winter and spring, depending on the severity

of the frontal systems associated with the winter rainfall

regime of the southwestern Cape. This harsh, icy environ-

ment gave rise to the specific epithet.

A subsequent visit in mid-December 1991 provided an

indication of the range of climatic extremes in which this

species survives, with very hot, dry conditions prevailing.

The steep-sided, rocky gullies in which this species is

found were, however, cooler, being shaded for much of

the day, and the humic soil was still damp. Figure 5 shows

the distribution of this species.

Vouchers: N. Barker 995. 996 (BOL, GRA, J, NBG. S, STE, UWC),

997 (BOL, K, PERTH, PRE), 1019 (BOL); Ellis 5620 (PRE).

INCERTAE SEDIS

A single unusual specimen, undoubtedly belonging to

the genus Pentameris (possessing an achene with apical

hairs) was collected from Cockscomb Peak by Ellis in 1987.

This specimen, Ellis 5605 (PRE), is morphologi-

cally similar in certain respects to P. glacialis, but differs

in its leaf anatomy.

ACKNOWLEDGEMENTS

I would like to thank Drs K. Balkwill, B. de Winter,

R.P Ellis, G.E. Gibbs Russell, E. A. Kellogg, D.J.B. Killick,

Bothalia 23,1 (1993)

45

FIGURE 14. Pentameris swartbergensis N.P. Barker. A, habit: erect culms (folded back, based on type material) and branched basal parts.

B-F, parts of spikelet and florets: B, glumes; C, whole basal floret; D, lemma (opened, flattened and viewed from inside of floret) showing

venation, lemma bristles, lemma lobes (adnate to bristles for most of their length) and geniculate awn; E, palea; F, developing fruit with

apical hairs, note that fruit appears to develop in a basipetal direction, uppermost region becoming swollen and associated surface sculptured,

appearing at apex first. A, X 0.5; B— E, X 4.8; F, X 14.6.

46

Bothalia 23,1 (1993)

FIGURE 15. — Pentameris glacialis

N.P. Barker. A, habit: show-

ing thin, decumbent culms; B,

indumentum of sheath, pubes-

cent along margin, sheath

mouth and leaf blade. C— H,

parts of s pikelet and florets: C,

glumes; D, whole basal floret

with mature fruit; E, lemma

(opened, flattened and viewed

from inside of floret) showing

venation, lemma bristles, lem-

ma lobes (adnate to bristles for

about half their length) and

geniculate awn; F, palea; G,

fruit with apical hairs; H,

spikelet. A, x 0.4; B— F, x

3.7; G, X 6; H, X 2.4.

O.A. Leistner, H.P. Linder and Mr E.R. Robinson for

their advice, discussions and comments on various aspects

of this work. Drs J. Rourke and O.A. Leistner are thanked

for assistance with the Latin diagnoses. In addition Dr Ellis

is acknowledged for the provision of his many micrographs

of leaf anatomy preparations and for comments on the

manuscript. The curators of the following herbaria are

thanked for providing access to, and/or loans of, their

specimens: B, BOL, GRA, SAAS, now housed in PRE,

K, NBG, OXF, P, PRE, S, SAM, STE and SAAS. I extend

my grateful thanks to Mrs Wilma Roux of the National

Botanical Institute for the illustrations of the new taxa,

Mrs S. Perold for assistance with the SEM and Mrs

A. Romanowski for the photographic work. I also acknow-

ledge the support of the National Botanical Institute, my

employer for the duration of this study. This work was un-

dertaken in partial fulfilment of an M.Sc. degree at the

University of the Witwatersrand.

REFERENCES

AMARASINGHE, V. & WATSON, L. 1988. Comparative ultrastruc-

ture of microhairs in grasses. Botanical Journal of the Linnean

Society 98: 303—319.

AMARASINGHE, V. & WATSON, L. 1989. Variation in salt secretory

activity of microhairs in grasses. Australian Journal of Plant

Physiology 16: 219-229.

BARKER, N.P. 1986. The shape and ultrastructure of the caryopsis of

Pentameris and Pseudopentameris species (Arundinoideae,

Poaceae). Bothalia 16: 65—69.

BARKER, N.P. 1989. The caryopsis surface of Pentameris and Pseudo-

pentameris (Arundinoideae, Poaceae) revisited. Bothalia 19:

134-136.

BARKER, N.P. 1990. The taxonomy of Pentameris Beau v. and Pseudo-

pentameris Conert. M.Sc. thesis, University of the Witwatersrand.

BARKER, N.P. in prep. a. The taxonomy of Pseudopentameris Conert

(Arundineae, Poaceae).

BARKER, N.P. in prep. b. Caryopsis surface features of southern African

Arundineae.

BURDET, H.M. 1976. Cartulae ad botanicorum graphicem. IX. Can-

dollea 31: 323-333.

Bothalia 23,1 (1993)

47

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

in southern Africa. In D. Meridith, The grasses and pastures

of South Africa. Central News Agency, Cape Town.

CLAYTON, W.D. & RENVOIZE, S.A. 1986. Genera graminum. Grasses

of the World. Her Majesty’s Stationery Office, London.

CONERT, H.J. & TUERPE, A.M. 1969. Karroochloa, eine neue Gattung

der Gramineen (PDaceae, Arundinoideae, Danthonieae). Sencken-

bergiana Biologica 50: 289—318.

DAVIDSE, G. 1988. A revision of the genus Prionanthium (Poaceae:

Arundinoideae). Bothalia 18: 143-153.

DESVAUX, N.A. 1831. Opuscules sur les sciences physiques et naturelles.

Pavie, Angers.

DE WET, J.M.J. 1956. Leaf anatomy and phylogeny in the tribe

Danthonieae. American Journal of Botany 43: 175-182.

DONOGHUE, M.J. & CANTINO, P.D. 1984. The logic and limitations

of the outgroup substitution approach to cladistic analysis.

Systematic Botany 9: 192—202.

DU PLESSIS, H. & SPIES, J.J. 1988. Chromosome studies on African

plants. 8. Bothalia 18: 119—122.

DURAND, T. & SCHINZ, H. 1895. Conspectus florae africae. Vol.

5. Friedlaender, Berlin; Klincksieck, Paris.

ELLIS, R.P. 1979. A procedure for standardizing comparative leaf

anatomy in the Poaceae. II . The epidermis as seen in surface view.

Bothalia 12 : 641-671.

ELLIS, R.P. 1980. Leaf anatomy of the South African Danthonieae

(Poaceae). III. Merxmuellera stricta. Bothalia 13: 191—198.

ELLIS, R.P. 1981. Leaf anatomy of the South African Danthonieae

(Poaceae). V. Merxmuellera macowanii, M. davyi and M. aureo-

cephala. Bothalia 13: 493—500.

ELLIS, R.P. 1985a. Leaf anatomy of the South African Danthonieae

(Poaceae). X. Pseudopentameris. Bothalia 15: 561-566.

ELLIS, R.P. 1985b. Leaf anatomy of the South African Danthonieae

(Poaceae). XI. Pentameris longiglumis and Pentameris sp. nov.

Bothalia 15: 567-571.

ELLIS, R.P. 1985c. Leaf anatomy of the South African Danthonieae

(Poaceae). XII. Pentameris thuarii. Bothalia 15: 573—578.

ELLIS, R.P. 1985d. Leaf anatomy of the South African Danthonieae

(Poaceae). XIII. Pentameris macrocalycina and P. obtusifolia.

Bothalia 15 : 579 -585.

ELLIS, R.P. 1986. Leaf anatomy of the South African Danthonieae (Poa-

ceae). XIV. Pentameris dregeana. Bothalia 16: 235—241.

ELLIS, R.P. & LINDER, H.P. 1992. Atlas of the leaf anatomy in Pen-

taschistis (Arundineae: Poaceae). Memoirs of the Botanical Sur\’ey

of South Africa No. 60.

GIBBS RUSSELL, G.E., REID, C., VAN ROOY, J. & SMOOK, L.

1985. List of species of southern African plants, edn 2, part 1.

Memoirs of the Botanical Survey of South Africa No. 51.

GIBBS RUSSELL, G.E., WATSON, L , KOEKEMOER, M., SMOOK,

L., BARKER, N.P., ANDERSON, H.M. & DALLWITZ, M.J.

1990. Grasses of southern Africa. Memoirs of the Botanical Survey

of South Africa No. 58.

GOLDBLATT, P. 1978. An analysis of the flora of southern Africa: its

characteristics, relationships and origins. Annals of the Missouri

Botanical Garden 65: 369—436.

GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern

Africa. Balkema, Cape Town.

HALL, A.V. & VELDHUIS, H.A. 1985. South African red data book:

plants— fynbos and karoo biomes. South African National

Scientific Programmes Report No. 117.

KARRON, J.D. 1987. The pollination ecology of co-occurring geographi-

cally restricted and widespread species of Astragalus (Fabaceae).

Biological Conservation 39: 179—193.

KUNTH, C.S. 1829. Revision des graminees. Paris.

KUNTH, C.S. 1833. Enumeratio plantarum I. Cotta, Stuttgart.

KUNTH, C.S. 1835. Enumeratio plantarum. Supplementum. Cotta,

Stuttgart.

LEHMANN, J.G.C. 1831. Pugillus plantarum III. Meissner, Hamburg.

LINDER, H.P. & ELLIS, R.P. 1990a. Vegetative morphology and

interfire survival strategies in the Cape fynbos grasses. Bothalia

20: 91-103.

LINDER, H.P. & ELLIS, R.P. 1990b. A revision of Pentaschistis

(Arundineae: Poaceae). Contributions from the Bolus Herbarium

12.

NEES AB ESENBECK, C.G. 1832. Plantae ecklonianae. Linnaea 7:

273 - 339.

NEES AB ESENBECK, C.G. 1841. Florae africanae australioris I.

Gramineae. Prausnitzianis, Glogau.

NORDENSTAM, B. 1980. The herbaria of Lehmann and Sonder in

Stockholm, with special reference to the Ecklon and Zeyher

collection. Taxon 29 : 279 —291.

PALISOT DE BEAUVOIS, A.M.F.J. 1812. Essai d’une nouvelle

agrostographie . Fain, Paris.

PHILLIPS, E.P. 1931. An introduction to the study of the South African

grasses with notes on their structure, distribution, cultivation,

etc. Central News Agency. South Africa.

PLATNICK, N.I. 1989. An empirical comparison of microcomputer

parsimony programs, II. Cladistics 5: 145-161.

RAB1NOWITZ, D. 1981. Seven forms of rarity. In H. Synge, The bio-

logical aspects of rare plant conservation : 205—217. Wiley, New

York.

RENVOIZE, S.A. 1981. The subfamily Arundinoideae and its position

in relation to a general classification of the Gramineae. Kew

Bulletin 36: 85-102.

RENVOIZE, S.A. 1986. A survey of leaf-blade anatomy in grasses VIII.

Arundinoideae. Kew Bulletin 41: 323-338.

ROEMER, J.J. & SCHULTES, J. A. 1817. Caroli a Linne equitis systema

vegetabilium 2. Cotta, Stuttgart.

ROHLF, F.J. 1988. NTSYS-pc. Numerical taxonomy and multivariate

analysis system. Exeter Publishing, New York.

SCHWEICKERDT, H ,G. 1938. Descriptions and notes on South African

grasses. Feddes Repertorium 43: 88-92.

SCHWEICKERDT, H.G. 1942. A taxonomic and anatomical study of

Elytrophorus Beauv. Annals of the Natal Museum 10: 191—214.

SPIES, J.J. & DU PLESSIS, H. 1988. Chromosome studies on African

plants. 6. Bothalia 18: 111-114.

SPIES, J.J., DU PLESSIS, H., BARKER, N.P. & VAN WYK, S.M.C.

1990. Cytogenetic studies in the genus Chaetobromus (Poaceae:

Arundineae). Genome 33: 646—658.

STAPF, O. 1897. Gramineae. In Flora capensis Vol. 7. Reeve, London.

STEUDEL, E.G. 1829. Bemerkungen ueber einige Cyperaceen und

Gramineen vom Vorgebirge der guten Hoffnung. Flora 12:

481-492.

STEUDEL, E.G. 1855. Synopsis plantarum graminearum. Metzler,

Stuttgart.

TOMLINSON, K.L. 1985. Comparative anatomical studies in Danthonia

sensu lato (Danthonieae: Poaceae). Aliso 11: 97—114.

.

Bothalia 23,1: 49-57 (1993)

Studies in the Marchantiales (Hepaticae) from southern Africa. 1. The

genus Dumortiera and D. hirsuta ; the genus Lunularia and L. cruciata

S.M. PEROLD*

Keywords: Dumortiera, D. hirsuta , Dumortieroideae, Hepaticae, Lunularia, L. cruciata. Lunula riaceae, Marchantiaceae, Marchantiales, taxonomy,

southern Africa, Wiesnerellaceae

ABSTRACT

The genera Dumortiera (Dumortieroideae, Marchantiaceae) and Lunularia (Lunulariaceae), are briefly discussed. Each

genus is represented in southern Africa by only one subcosmopolitan species, D. hirsuta (Swartz) Nees and L. cruciata

(L.) Dum. ex Lindberg respectively.

UITTREKSEL

Die genusse Dumortiera (Dumortieroideae, Marchantiaceae) en Lunularia (Lunulariaceae) word kortliks bespreek. In

suidelike Afrika word elke genus verteenwoordig deur slegs een halfkosmopolitiese spesie, D. hirsuta (Swartz) Nees en L.

cruciata (L.) Dum. ex Lindberg onderskeidelik.

DUMORTIERA Nees

Dumortiera Nees ab Esenbeck in Reinwardt, Blume

& Nees ab Esenbeck, Hepaticae Javanicae, Nova Acta

Academiae Caesareae Leopoldina-Carolinae Germanicae

Naturae Curiosorum XII: 410 (1824); Gottsche et al.\ 542

(1846); Schiffner: 35 (1893); Stephani: 222 (1899); Sim:

25 (1926); Muller: 394 (1951-1958); S. Amell: 52 (1963);

Hassel de Menendez: 182 (1963). Type species: Dumor-

tiera hirsuta (Swartz) Nees.

Synonymy according to Nelson & Parnell: 35 (1992):

Hygropyla Taylor: 390 (1836).

Hygrophila Taylor (orth. var.) in Mackay: 53 (1836) non

R. Br. (1810).

Hygrophyla Taylor (orth. var.) in Mackay: X (1836).

Hygropila Taylor (orth. var.) in J.D. Hooker & T. Taylor:

576 (1844).

Askepos Griffith 2: 340 (1849).

Thallus, large, flat and thin, dark green, in overlying

patches; on shaded, damp soil or on wet rocks. Branches

with apical innovations or dichotomously furcate, occa-

sionally lateral, moderately divergent, thickened over

midrib, wings gradually thinning toward margins; apex

emarginate.

Dorsal epidermis very thin-walled, temporary, vestigial,

air pores absent, air chambers mostly reduced to 1- or

2-celled vestiges, reported rarely to have reduced chloro-

phyllose filaments, cells containing numerous fairly large

chloroplasts; storage tissue compact, confined to ventral

part of midrib; oil cells rare, each with a single, large oil

body; rhizoids along midrib and ventrally appressed

against wings, mostly smooth, occasionally tuberculate;

scales ventral over midrib, vestigial and evanescent.

* National Botanical Institute, Private Bag X101, Pretoria 0001.

MS. received: 1992-03-31.

Monoicous or dioicous. Antheridia sunken in subses-

sile disciform receptacles, which are fringed with bristles

and borne singly at apex of thallus on short bifurrowed

stalk. Archegonia in groups of 8—16 in saccate, fleshy

involucres, on lower surface of 6-8-lobed disciform

receptacle with marginal sinuses dorsally, raised on stalk

with two rhizoidal furrows; after fertilization and

maturation, each involucre generally containing a single

sporophyte consisting of foot, seta and capsule; capsule

wall unistratose, with annular thickenings, dehiscing irre-

gularly. Spores small, papillose. Elaters slender, tapering,

1— 3-spirate. Gemmae absent.

Dumortiera hirsuta (Swartz) Nees in Reinwardt,

Blume & Nees ab Esenbeck, Nova Acta Academiae

Caesareae Leopoldino-Carolinae Germanicae Naturae

Curiosorum XII: 410 (1824); Gottsche et al.\ 542 (1846);

Spruce: 566 (1885); Stephani: 224 (1899); Macvicar: 41

(1926); Sim: 25 (1926); Muller: 396 (1951); S. Amell: 52

(1963); Hassel de Menendez: 182 (1963). Type: Jamaica,

leg. Swartz s.n. [S, holo.!; MW, iso. (Hb. Hoffm. No.

8497); UPS, fide Grolle 1976],

Marchantia hirsuta Swartz: 145 (1788).

M. irrigua Wilson ex Hooker in Smith: 106 (1833). Hygropyla irrigua

(Wilson) Taylor: 390 (1836); Mackay: 54 (1836); Stephani: 150 (1899).

Dumortiera irrigua (Wilson) Nees: 159 (1838). D. hirsuta var. irrigua

(Taylor) Spruce: 566 (1885). Type: Ireland, Tbrk Cascade, near Killamey,

Mr Wilson.

D. hirsuta var. angustior Gottsche, Lindenberg & Nees: 544 (1846).

D. hirsuta var. intermedia Gottsche, Lindenberg & Nees: 544 (1846).

D. velutina Schiffner: 256 (1893).

Askepos brevipes Griffith: 340 (1849). Type: India, in sylvis umbro-

sis Tingrei agri, II 1836.

Thallus hygrophyllous, large and creeping, broadly and

rather irregularly strap-shaped, uniformly dark green,

translucent, thin and flat, but often marginally undulate,

wet (Figure 1A); crisped and shrivelled, dull, unable to

survive dry; in crowded, overlying patches, once to several

times dichotomously or occasionally laterally furcate or

50

Bothalia 23,1 (1993)

FIGURE 1. — Dumortiera hirsuta. A, dorsal view of thallus; B, ventral view of thallus; C, male plant with disciform receptacles at apex; D, young

female receptacle seen from above; E, young female receptacle seen from below; F, transverse section of thallus; G, margin of thallus

with hairs; H, transverse section of thallus, much enlarged; I, lower cells of costa and vestigial scales in transverse section; J, filiform

scale from top of stalk; K, transverse section of stalk with two rhizoidal furrows; L, capsule wall cells with thickenings; M, transverse

section of capsule wall. A, F-H, S.M. Perold2694: B, 5. M. Perold2634: C, Nicholas 1176: D, E, I, H. Anderson CH 13495: J-L, Doidge

CH 3581. Scale bars: A— C = 2 mm; D— F, J = 1 mm; G, K = 100 jxm; H, I, L, M = 50 /xm. Illustrations by G. Condy.

Bothalia 23,1 (1993)

51

with apical innovations; branches 50-95 x 8— 13(— 22)

mm and ± 500 /tm thick over median, ventrally bulging

midrib (also visible from above), laterally gradually

thinning out into wide wings (Figure IF); apex shortly

emarginate; margins undulate, occasionally scalloped,

sparsely hirsute, hairs ± 250.0 /xm long, ± 12.5 /tm wide

at base, tapering slightly to somewhat blunt tip (Figure

1G).

Epidermis, pores and air chambers absent; dorsal cells

from above, 4—6-sided, variable in shape and size, 27-57

x (17-)22-32 /xm, in cross section ± 15 /on thick,

containing chloroplasts; marginal cells thin-walled, long-

or short-rectangular, 25-62 x 12-20 /im; midrib with

+ 18 rows of cells of which upper 3 rows larger, rounded,

up to 100 x 137 /xm, with somewhat wavy walls and

scattered chloroplasts, lower group of cells angular,

smaller, ± 30 /im wide, lacking chloroplasts; wings

medianly with 3 rows of large cells, decreasing in number

to one row at margin, covered by dorsal and ventral layer

of smaller chlorophyllose cells; oil bodies quite rare, for

the most part confined to scattered cells in the midrib,

yellow-brown, elongate or round, ± 40 /xm across; rhi-

zoids below midrib numerous, mostly smooth, 15-20 /xm

wide, occasionally tuberculate, 5 /xm wide, also in widely

spaced, arching strands below wings, radiating toward

margins and appressed to ventral face of wings (Figure

IB) or scattered; ventral scales hyaline, vestigial (Figure

II), without appendages, evanescent, only near apex.

Monoicous or dioicous. Antheridiophore subsessile, dis-

ciform, 2.75 mm in diameter, 0.6 mm thick in centre, flat-

tening toward sides, containing antheridia, 470 x 250 /im,

± ovate, acropetally arranged, but not in radiating rows,

sunken into disc, borne singly at apical notch of thallus

(Figures 1C; 2A) on very short, 1 mm diameter,

bifurrowed stalk, with rhizoids lining furrows; hyaline, fili-

form scales on ventral face of disc, the latter encircled by

dense outer fringe of bristle-like hairs, 600-1100 pm long,

base 20—25 /xm wide, a few scattered hairs dorsally. Ar-

chegoniophore disciform, 3.75 mm in diameter; dorsal face

(Figure ID) becoming radially grooved by sinuses; ven-

tral face 6— 8-lobed (Figure IE) in radiating rows and then

almost star-shaped, sparingly bristled; archegonia

in groups of 8-16, enclosed in green, fleshy, saccate

involucres, long archegonial necks protruding through

narrow slit-like apical openings; receptacle eventually

raised on stalk arising at apical notch of thallus between

overlapping sides, + 40 mm long and ± 925 /xm wide,

cortical cells + 15.0 x 12.5 /xm, inner cells + 50 x 30

/xm; along its length 2 furrows lined with rhizoids (Figure

IK) and its top (where joined to disc) encircled by nume-

rous filiform, hyaline scales (Figure U), up to 3750 /xm

long and 300 /xm wide at base, apex only ± 3 cells wide,

cells mostly ± 125 x 22 /xm; at maturity generally only

one sporangium borne in each involucre, rarely two and

quite often none, elliptical, 2.5 x 1.5 mm, supported on

elongating seta and protruding from membranous calyp-

tra, pseudoperianth lacking; capsule wall unistratose

(Figure 1M) with annular or semi-annular thickenings

(Figure 1L), dehiscing irregularly via longitudinal suture

lines and releasing large numbers of elaters and spores

(Figure 2B). Spores golden brown, semi-transparent, distal

face rounded, proximal face ± flat to slightly peaked in

the centre, triradiate mark indistinct, 25—30 /xm across

its longer axis, ornamented with numerous nodules or

tubercles, irregular in size and shape (Figure 3A). Elaters

yellow-brown or orange-brown, mostly with doubly spiral

strands (Figure 3B), ends tapering, 225 —470 x 7.5—10.0

/xm in middle and 3.5 /xm wide at tips. Gemmae absent.

Chromosome number, n = 9 (Berrie 1960, Bomefeld

1987); n = 18 (Tatuno 1941); n = 27 (Tatuno 1941, Berrie

1958). According to Tatuno (1938, 1939) the 9, 18 and 27

chromosome sets present different races with different

distributions and edaphic ecology, but he considers them

as belonging to the same species.

Only a few of the specimens examined were fertile; most

of these were dioicous, with male and female receptacles

borne on separate plants. Monoicous plants were quite

rarely found, but have frequently been reported in the

literature and even bisexual receptacles are known (Evans

1919).

All southern African specimens have been identified as

belonging to D. hirsuta : the dorsal face of the thallus is

always + smooth and lacks papilliform cells or traces of

air chambers. D. nepalensis and D. velutina, which our

material resembles, are considered to belong to the poly-

morphic species D. hirsuta (Grolle & Piippo 1984). Amell

(1963) referred to material from southern Africa (Cape

Province, Transvaal, Natal) as D. hirsuta var. nepa-

lensis. Schuster (1992) considers D. nepalensis (Taylor)

Nees to be a distinct taxon under the name of D. hirsuta

subsp. nepalensis (Taylor) Schuster which is not found in

Africa. D. hirsuta is subcosmopolitan and is widespread

in tropical and temperate regions, generally growing in

sheltered, wooded, shaded and damp areas, i.e. it is hygro-

philous. In southern Africa it is known from northern and

eastern Transvaal, Swaziland, Natal, Zululand and eastern,

southern and southwestern Cape (Figure 4). Further north-

wards in Africa, D. hirsuta is reported from (or has been

collected in) Zimbabwe (Best 1990), Zambia [S.M. Perold

2670 (PRE)], East African Mountains (Amell 1956), East

Africa (Bizot etal. 1985), Tanzania, Malawi, Mozambique

(Bizot & Poes 1979), Rwanda and Burundi (Vana et al.

1979).

On the basis of flavonoid data, Campbell et al. (1979)

found chemical affinities between Dumortiera and Wies-

nerella, suggesting that they belong in a common family,

Wiesnerellaceae (Inoue 1976). This was accepted by Grolle

(1983), but Schuster (1984) has retained Dumortiera in the

Marchantiaceae, creating a new subfamily, Dumortie-

roideae Schust., for it and designating Dumortiera as the

type genus. Dumortiera has also been classified in the

Dumortieroideae by Bischler (1988), who regards Dumor-

tiera and Wiesnerella as morphologically remotely related,

although having similar flavonoid patterns. The treatment

of Schuster (1984) is followed here, even though the

phylogenetic position of Dumortiera remains a matter of

some controversy, as conceded by Schuster (1984). Sup-

porting the above treatment are terpene studies of the

genera by Asakawa et al. (1979, 1980a, 1980b, 1981), who

found that Wiesnerella denudata and Conocephalum coni-

cum shared 17 of these terpene compounds, but only had

three in common with Dumortiera hirsuta. Luteolin 5-0-

glucuronide, the flavonoid shared by Dumortiera and Wies-

nerella which was considered diagnostic for the Wiesnerel-

laceae by Campbell et al. (1979), has since also been

52

Bothalia 23,1 (1993)

FIGURE 2. — Dumortiera hirsuta. A,

male plant with subsessile

antheridiophore from above; B,

stalked archegoniophore with

dehisced sporangia, front one

releasing many spores and

elaters. A, B, Koekemoer 989 ,

x 9.

reported for Conocephalum by Porter (1981). Whittemore

(1991) therefore cautions against drawing taxonomic con-

clusions from a small number of compounds.

SPECIMENS EXAMINED

TRANSVAAL. — 2230 (Messina): Entabene, (— CC), Bottomley (PRE);

Entabene, (— CC), Schelpe 6020 (BOL); Soutpansberg, Entabene, (— CC),

Thomas 856 (PRE). 2329 (Pietersburg): Pietersburg, (—CD), Van Vuuren

1469 (PRE); Haenertsburg, ( — DD), Putterill 3604 (PRE). 2330 (Tza-

neen): Woodbush For. Res., Magoebaskloof, near stream, (— CC), H.

Anderson CH 13495, CH 13499 (PRE); Bosnian 3188 , (PRE); De Hoek

For. Res., Debengeni Falls, on stream bank, beyond foot bridge, (— CC),

S.M. Perold 2634 (PRE); Woodbush, locally abundant in deep shade

on streambank, (— CC), Schelpe 6070 (BOL). 2331 (Phalaborwa): Letaba,

(-DC), Scheepers 984 (PRE). 2430 (Pilgrim’s Rest): Farm Cyprus near

Ofcolaco, in gorge named ‘Terrible Hollow’, (— AB), H. Anderson CH

4527 (PRE); Mariepskop Forestry Water Works, on outside of water-

tank, also under overhang, dense shade, (— DB), Vorsier 87C (PRE);

Mariepskop, near dam in KJaserie River, montane forest, on rock against

bank next to stream, in shade, (— DB), Vorster 572; Mariepskop For.,

Bedford footpath in forest on soil of streambed, full shade, (— DB), Vor-

ster 1398 (PRE); Mariepskop For., Blyde River footpath, on exposed

tree root at streamside in forest shade, (— DB), Vorster 1472 (PRE);

Mariepskop, Blyde River footpath , hanging from vertical sandstone rocks

in forest shade, (-DB), Vorster 1473 (PRE); Mariepskop, Magalieskop

Res. For., on damp earth bank, dense shade, (— DB), Vorster 1815 (PRE);

Mariepskop, ( — DB), Van der Schijff 4482 (PRE); Mariepskop, Klaserie

River by dam, beneath rocks, (— DB), Van der Schijff 6291 (PRE); Mount

Sheba Nat. Res., at the ‘Grotto’, on dripping cliffs, forming extended

mats on wet vertical rock face, (—DC), Jacobsen 4421 (PRE); Mount

Sheba Nature Reserve at the ‘Grotto’, on dripping rock cliffs, (—DC),

Perold & Koekemoer 2864 (PRE); Pilgrim's Rest, (— DD), Van der Schijff

6367 (PRE). 2527 (Rustenburg): Nature Res. Cederberg kloof, near Uto-

pia, (— CA), Koekemoer 972 (PRE). 2530 (Lydenburg): Farm Klipsteen,

between Lydenburg and Dullstroom, at waterfall, on rock, (-AB), H.

Anderson CH 13446 p.p. (PRE); Sabie, Lone Creek Falls, on soil near

footpath, (—BA), S.M. Perold 2694 (PRE); Sabie Gorge, (— BB), V.A.

Wager 21 (PRE); on road to Lydenburg, at turnoff to Witklip, Coromandel

Farm, at waterfall, (-AB), Perold & Koekemoer 2839, 2844 (PRE); Nel-

spruit Dist., Rooiwal, (— BC), Bosnian 3180 ( PRE); Rosehaugh, ( — BD),

T.R. Sim CH 1286 (PRE); Kaapsehoop, (— DB), H.A. Wager 47 ( PRE);

Berlin State Forest, Kaapsehoop hiking trail near Battery Creek, (-DA),

Koekemoer 973, 975 (PRE).

Bothalia 23,1 (1993)

53

FIGURE 3. — Dumortiera hirsuta. A, spore, distal view; B, part

of elater. A, B, H. Anderson CH 4527. A, x 1730; B,

x 480. SEM micrographs by S.M. Perold.

SWAZILAND. —2531 (Komatipoort): King Forest, Havelock, occa-

sionally on wet shaded earth bank in forest, (— CC), Schelpe 6195 (BOL).

NATAL.— 2731 (Louwsburg): Ngoma For., bank of river, (—CD),

Gerstner 4386 (PRE); Ngoma For., Cetshwayo waterfall walk, vertical

slope, seepage, in deep shade, (—CD), Glen 2881 (PRE); Ngoma For.,

along path in Ntendeka, alongside streams in forest interior, (—CD),

Nicholas 1176 (PRE); Ngoma Forest, occasionally on deeply shaded earth

banks in forest, (—CD), Schelpe 6237 (BOL); Ngoma For., (—CD), T.R.

Sim CH 1302 (PRE); Ngoma For., Ntendeka, in forest, (-CD), A.E.

van Wyk 6973 (PRE). 2828 (Bethlehem): Mont-aux-sources, (— DD),

Doidge 171 (BOL, PRE). 2831 (Nkandla): Nkandla Forest, common on

rocks in and near water in shade, (— CA), Nixon 57 (BOL); Eshowe,

Signal Hill, (—CD), Van der Plank CH 1290, CH 1298 (PRE); Ngoya,

(-DC), T.R. Sim CH 1289 (PRE). 2929 (Underberg): Mpendle Dist.,

6 miles along Everglades/Boston Road River on stream bank, in shade,

(— DB), Moll 726 (BOL, PRE); Home Rule, Polela, (-DC), T.R. Sim

CH 1305 (PRE); Xumeni For., (— DD), Doidge 3581 (PRE); Donnybrook,

(-DD), Scott 3756 (PRE). 2930 (Pietermaritzburg): Karkloof, (—AC),

T.R. Sim CH1291 (PRE); Buccleugh, (—AD), T.R. Sim CH 1301 (PRE);

Pietermaritzburg, Town Bush, (— CB), T.R. Sim 7523, 7534; Waterfall,

(— CB), T.R. Sim 7548, 7574, 7586 (PRE); Pietermaritzburg, (-CB),

T.R. Sim 1296 (PRE); Sweetwater stream, (— CB), T.R. Sim 1300 (PRE);

Zwaartkop, (— CB), T.R. Sim CH 1328 (PRE).

CAPE.— 3226 (Fort Beaufort): Katberg For., (-BC), Garabedian

S. A.M.H. 49743 (BOL; PRE); Hogsback, (-DB), T.R. Sim 1895; Van

der Bijl 176; Young 1303 (PRE). 3 227 (Stutterheim): Perie Forest, (-CC),

T. R. Sim 7534 (BOL). 3318 (Cape Town): Cape Town, (-CD), H.A.

Wager 28 (PRE). 3322 (Oudtshoom): Rust en Vrede, at Oudtshoom,

(— CA), ex Herb. C. vanden Berghen (BOL). 3323 (Willowmore): Gou-

na Forest, near Lily vlei, (— CC), S. Amell 1736 (BOL); Bloukrans River

Pass, at bridge in forest, on rocks along stream, (—DC), Stirton 9648

(PRE); Bloukrans, along pass in river ravine, on vertical cliff face,

(-DC), Zantovska 155 (PRE). 3419 (Caledon): Oudebos, Rivier-sonder-

end, (— BB), Thome 3617 (BOL, PRE). 3420 (Bredasdorp): Leeuwrivier

Mountains, Swellendam, (— AB), Stokoe 9486 (BOL). 3423 (Knysna):

Knysna, Garden of Eden, (— AA), S. Amell 2096 (BOL); Knysna,

(— AA), T.R. Sim 1292 (PRE).

LUNULA RI A Adanson

Lunularia Adanson , Families des plantes 2: 15 (1763);

Micheli: 4 (1729); Nees ab Esenbeck: 29 (1838); Gottsche

et al.\ 510 (1846); Schiffner: 35 (1893); Stephani; 216

(1899); Howe: 59 (1899); Macvicar: 38 (1926); Sim 123

(1926); Muller: 366 (1951-1958); S. Amell: 73 (1963);

Hassel de Menendez: 125 (1963). Type species: Lunularia

cruciata (L.) Dum. ex Lindb.

Selenia J. Hill: 120 (1773) nom. illeg.

Staurophora Willd. 3: 101 (1809).

Dichominum Neck.: 345 (1790) (as subgenus).

Marsilia O. Kuntze 2: 837 (1891).

Sedgwickia S. Bowdich: 35 (1825).

Thallus large, flat, somewhat glossy, green, in overlying

patches, sometimes in extensive turfs; on damp soil in old

gardens, nurseries and forested areas, may have been

introduced into southern Africa. Branches dichotomously

furcate toward apex, new growth by apical or lateral

innovations; thickened over midrib, gradually thinning

toward slightly undulate, hyaline margins; apex emar-

ginate. Dorsal epidermis persistent, hyaline, cell walls

sometimes thickened at comers or entirely; air pores

simple, elevated and conspicuous, surrounded by several

concentric rings of cells, leading below into individual air

chambers, these in one layer and floored by chlorophyl-

lose layer of 3— 5-celled erect, branched filaments; storage

tissue compact, cells colourless, sometimes with pitted

walls; scattered cells throughout with single, large, brown

oil body; rhizoids numerous, between ventral scales and

on midrib, some smooth, others tuberculate; scales ventral,

in curved parallel rows on either side of midrib, with round

or reniform appendages, several cells con-

taining oil bodies.

Dioicous. Antheridia sunken into slightly elevated, oval

or kidney-shaped, disciform receptacles encircled by

raised, membranous sheath, at sides of male thalli

(although originally terminal). Archegonia enclosed in

terminal receptacles, but by continued growth of thallus

FIGURE 4. — Distribution of Dumortiera hirsuta, #; and Lunularia

cruciata, □, in southern Africa.

54

Bothalia 23,1 (1993)

FIGURE 5 .—Lunularia cruciata. A, dorsal view of thallus with gemma cups; B, female plant with young archegoniophores; C, ventral view

of thallus with new lateral branch; D, transverse section of thallus; E, transverse section of midrib region, much enlarged; F, transverse

section of air chamber; G, air pore seen from above; H, margin of thallus with hyaline cells, seen from above; I, older scale; J, young

scale; K, longitudinal section through young archegoniophore; L, longitudinal section through gemma cup; M, gemma. A, D— H, J,

L, M, S.M. Perold 2821\ B, C, S.M. Perold 1996. Scale bars: A-C = 2 mm; D, E, K, L = 1 mm; I, J = 500 /«n; F, M = 100 ^m;

G, H = 50 /im. Illustrations by A. Pienaar. Figure IK partly after Benson-Evans & Hughes fig. 4.

Bothalia 23,1 (1993)

55

laterally situated, conical, white and bud-like when young,

sheathed in layers of scales and slightly sunken into

rounded depression with distinct rim; after fertilization

and further growth, four tubular involucres in the form

of a cross are formed, each enclosing one or two sporo-

phytes, composed of foot, seta and capsule, raised on

unfurrowed, hairy stalk; capsules eventually exposed by

elongation of seta, the wall unistratose, lacking annular

thickenings, dehiscing by 4 valves. Spores very small,

green or brown, smooth. Elaters long, tapering and

bispiral. Gemmae numerous, disc-shaped, inside crescent-

shaped ridge; wholly diagnostic. Sporophyte virtually

unknown in southern Africa.

Lunularia cruciate (L.) Dum. ex Lindberg, Notiser

Sallskap pro Fauna et Flora Fennica Forhandlingar 9: 298

(1868); Howe: 60 (1899); Macvicar: 40 (1926); Sim: 24

(1926); Muller: 366 (1951-1958); S. Amell: 73 (1963);

Hassel de Menendez: 126 (1963); E.O. Campbell: 31

(1965). Type: In Europae umbrosis [OXF, syn.; H-SOL,

isosyn., fide Grolle (1976)]. For detailed synonymy see K.

Muller (1951-1958).

Thallus moderately large and flat, ribbon-like or margins

somewhat irregular (Figure 5A), glossy, bright green to

yellowish green, with outlines of subdorsal air chambers

faintly visible from above, each of the polygonal areas with

a central air pore, when wet; reticulum indistinct, leathery,

when dry; in crowded overlying patches, dichotomously

or irregularly furcate or with apical or lateral innovations

from ventral side of thallus (Figure 5C). Branches 40—55

x 5 — 8( — 10) mm, ± 650( - 1000 ) pm thick over median,

ventrally bulging midrib, gradually thinning out laterally

into wide wings (Figure 5D); apex emarginate or sinusoi-

dal; margins slightly undulate and somewhat scalloped,

with outer 4(5) cell rows hyaline.

Dorsal epidermal cells hyaline, in one layer, 5- or

6-sided to irregular in shape, 35—50 x 20—30 pirn, thin-

walled to somewhat thicker-walled or only thickened at

corners, in cross section 20—25 pm thick; marginal cells

with outermost row short- to long-rectangular (Figure 5H),

15-27 x 10-12 pm, cells of inner rows polygonal, 17-27

x 25 pm\ air pores simple, oval, raised, 17—25 x 12—20

pm in diameter, bordered by 3—5 rings of curved, smaller,

thin-walled cells (Figure 5G), 7—12 x 17—27 pm, outer

row of cells somewhat larger, 15—20 x 20—22 pun; air

chambers with domed roof (Figure 5F) raised 37—52 pun

above filaments, laterally separated by non-chlorophyllose

1 unistratose partitions obscured by filaments, floored by

dense chlorophyllose layer, ± 70 pirn thick, of 3— 5-celled

erect, branched filaments, filled with chloroplasts, top cell

often clavate, + 20 x 15 pun, others 20 x 12 pm; midrib

below assimilation cells, with ± 15(— 20) rows of com-

pact colourless storage cells (Figure 5E), 50—75 x 45 —50

pun, becoming smaller ventrally, some with pitted walls,

layers gradually decreasing in the wings; scattered cells

with brown oil bodies, round or oval, 30 x 30—45 x 27

pan; ventral epidermal cells 30—40 pirn wide, 15—25 pirn

! thick in cross section; rhizoids on midrib between scales,

numerous, smooth, ± 30 pirn wide, tuberculate, (10—)

17—22 pirn wide. Scales hyaline, some basal cells purple,

on either side of midrib, stretched across ventral face of

wings, near apex of thallus, base ± 1375 x 550 pun, with

rounded, constricted appendage (Figure 5J), ± 400 x 250

pun; cells mostly polygonal, ± 65 x 25 pun, some smaller,

with oil body 22 x 25 pirn, almost entirely filling cell;

older scales larger (Figure 51), base up to 1000 x 4250

pirn, appendage ± 600 pirn wide.

Dioicous. Male plants quite rare in southern Africa.

Anther idiophore on alternate sides, having originally

developed in terminal sinus near apex (Figure 6A), slighdy

raised, ovate, flatfish discs, 2—3 mm in diameter, encircled

by membranous sheath with crenate edges, containing

numerous antheridia, individually sunken in flask-shaped

cavities (Figure 6B) , opening above by pores. Archegonio-

phore originally also developing in terminal sinus near

apex, with further growth of thallus leaving it behind, so

that it appears lateral in position on alternate sides of

thallus (Figure 5B); commences as domed disc ± 200 x

375 pun, bearing several archegonia in radiating rows and

attached by very short stalk to floor of shallow, round hol-

low, 1100 pun wide x 300 pun deep, with crenate rim;

sheathed in ± 3 layers of scales (Figure 5K), outer layer

formed by fused, shaped scales, bulging in centre and con-

stricted below and above, upper edge irregularly fringed

with filiform cellular appendages of up to 7 rectangular

cells, ± 35 x 15 pirn, cells in body of scale thick-walled,

5- or 6-sided, up to 57 x 25 pun, in between numerous

smaller cells, 4- or 5-sided, 25 x 15 pirn, almost entirely

filled with single oil body; between scales and especially

from base of archegoniophore and arching over it,

numerous uniseriate, long hairs of 16—20 cells each.

Further development not recorded as fertilization did

not take place, nor are sporophytes available for study.

Gemmae numerous, disc-shaped (Figure 5M), notched

FIGURE 6. —Lunularia cruciata . A, antheridiophore seen from above;

B, longitudinal section through antheridiophore, with one flask-

shaped antheridium in place. A, Wilman BOL. No. 24870: B.

Koekemoer 1004. A, x 22; B, X 38. A, SEM micrograph; B,

LM photograph.

56

Bothalia 23,1 (1993)

by 2 opposite, lateral growing points, ± 430 in

diameter when mature, developing upright on short stalk,

inside crescent-shaped cupule (Figure 5L), ± 3 mm wide

with crenate to entire ridge on proximal side. Chromo-

some number n = 8 (Heitz 1927); n = 9 = 8 + x/y2

(Lorbeer 1934); n = 9 (Bornefeld 1987).

DISCUSSION

As mentioned in the description, male plants with

antheridial receptacles are exceedingly rare in southern

Africa and the only two, Wilman BOL No. 24870, on loan

from BOL and Koekemoer 1004 (PRE), were received after

completion of Figure 5 and therefore not illustrated there

(see Figure 6). Male plants are so rarely seen, that Sim

(1926) had categorically stated that they are not present

in South Africa, but then added ‘so far as is known’. He

must have been unaware of Saxton’s find. No female plants

with fertilized archegonia and mature sporophytes were

available for study; even plants with young archegonio-

phores are quite rare, judging both from personal

experience and from the literature (Saxton 1931; Good-

man 1956; E.O. Campbell 1965). Saxton (1931) found both

male and female plants in Cape Town in 1908, but had

to wait for almost 20 years for plants with mature

archegoniophores (forwarded from Dartmouth, England)

to complete his studies on the life history of L. cruciata.

Giffen sent plants with young archegoniophores from

Oranjezicht, Cape Town to Sim (Sim 1926) and Auret (as

mentioned in Benson-Evans & Hughes 1954) reported the

regular production of female branches in the vicinity of

Johannesburg. Of my own collections, only S.M. Perold

1996 from Devon Valley Hotel, Stellenbosch (November

1987), had young archegoniophores.

Since so many collections are from nurseries or city

gardens, it would appear that L. cruciata may have been

introduced into southern Africa. It is not frequently found

here, most collections being from the southwestern Cape,

a few from southern Transvaal and some from Natal, and

then quite frequently from nurseries (Figure 4). Further

north in Africa, L. cruciata is known from Zimbabwe

(Best 1990), Malawi (Nyika Plateau, S.M. Perold 2667,

2676 (PRE)); East African Mountains (Arnell 1956);

Tanzania (Serengeti) (Vanden Berghen 1965); Uluguru

Mountains, Rungwe Mountains (Bizot & Poes 1979);

Congo Rep. (Katanga) (Vanden Berghen 1965). Poes (pers.

comm.) states that in East Africa, L. cruciata is found

in many natural habitats in the montane forest belt, mostly

on young volcanoes such as Mt Elgon and Mt Meru and

even on the Comoro Islands.

Lunularia cruciata is quite widespread in the southern

hemisphere but regarded by Engel & Schuster (1982) as

probably Laurasian in origin. Since sexual reproduction

and subsequent spore production are so rare, its gemmae

obviously present a highly effective means of ensuring its

dispersal, which is most likely aided by human activities.

On the basis of its flavonoid chemistry, Campbell et al.

(1979), include it in the Marchantiaceae, but in the present

treatment Grolle (1983) is followed and L. cruciata is

classified in the monotypic family, Lunulariaceae Klinggr.

(1858).

Its phylogenetic position seems rather unclear, Schuster

(1984b) arguing that he would place it ‘low’ in the

Marchantiales on account of its high level of seta re-

tention, 2—3 sporophytes per gynoecium and a capsule

with 4 well-defined valves. On the other hand, he expresses

the viewpoint (Schuster 1984b) that its archegonio-

phore, clearly formed from two dichotomies, each

producing archegonia, is complex and therefore an

advanced feature.

SPECIMENS EXAMINED

TRANSVAAL.— 2528 (Pretoria): Pretoria, Union Buildings Nurseries,

(— CA), Bottomley CH 135 (PRE); Pretoria, National Botanical Institute

Nurseries, (— CA), S.M. Perold 2821 (PRE). 2627 (Potchefstroom),

Vereeniging, ( — DB), T.R. Sim CH 1283 (PRE); Roodepoort, Sterlig

Nursery, on gravel and brick walls of flowerbeds, (— DD), Koekemoer

1004 (PRE).

NATAL.— 2930 (Pietermaritzburg): Hilton Road, (— CB), T.R. Sim

CH 1279, CH 1280 (PRE). 2931 (Stanger): Durban, Silverglen Nurser-

ies, (- CC), S.M. Perold 2805 (PRE).

CAPE. — 3318 (Cape Town): Kirstenbosch, (—CD), S. Arnell 412

(BOL); Window Gorge, Table Mountain, (—CD), S. Arnell 405 (BOL);

Claremont Park, (—CD), Garside 6653 (BOL); near Round House,

(—CD), Garside 6132 (BOL); Oranjezicht, (—CD), Giffen CH 1281

(PRE); Glen Picnic Resort, just below Round House, Lion’s Head,

(—CD), S.M. Perold 645 (PRE); Round House, Lion’s Head, on soil

beneath trees, (—CD), S.M. Perold 650 (PRE); Newlands Forest, on

soil under trees, (—CD), S.M. Perold 662 (PRE); Cape Town, (—CD),

T.R. Sim CH 1278, CH 1282 (PRE); Skeleton Gorge, rock face, very

wet forest, (—CD), Stirton 9415 (PRE); mountain slopes above Kirsten-

bosch, (—CD), Stokoe s.n. (BOL); Kirstenbosch, (—CD), Wilman 24870

(BOL); Stellenbosch, (— DD), Burtt Davy CH 1277 (PRE); Pniel, on

tree trunk mixed with moss, (— DD), Morley 314 (PRE); Stellenbosch,

Devon Valley Hotel at garden fence, on soil, (— DD), S.M. Perold 1990

(PRE); Stellenbosch, Devon Valley Hotel, on soil bank behind hotel,

(— DD), S.M. Perold 1996 (PRE); Stellenbosch, (-DD), T.R. Sim CH

1284 (PRE); S Paarl, Landskroon, in kloof, (— DD), Volk 81/071 (BOL,

PRE). 3319 (Worcester): W of Franschhoek, Waterval Farm, on earth

bank, ditch next to dirt road, (— CC), S.M. Perold 633 (PRE); 4 km

N of Villiersdorp, Elandsriver Road, Du Toitsberge, near waterfall on

Sneeukop, (—CD), S.M. Perold 623 (PRE). 3322 (Oudtshoom): Ge-

orge, NE of Hawthomdene Hotel , at roadside on side of earth water fur-

row, (—CD), S.M. Perold 920 (PRE). 3418 (Simonstown): Constantia,

on soil on damp rock face, (— AB), S.M. Perold 656 (PRE).

ACKNOWLEDGEMENTS

I wish to express my gratitude to Dr R. Grolle for his

information regarding the publication date of ‘Hepaticae

Javanicae’ by Reinwardt et al. , as well as for drawing my

attention to Nelson & Parnell’s publication on the

synonymy of the various forms of Hygropyla etc. under

Dumortiera. I also thank Dr Grolle, Prof. T. Poes and

Prof. O.H. Volk for critically reading the manuscript and

for their valuable suggestions. Further thanks are due to

the curator of BOL for the loan of specimens; to the artists,

Ms G. Condy and Ms A. Pienaar; to the typist, Mrs J.

Mulvenna and to the photographer, Mrs A. Romanowski.

Also to my colleagues at NBI, particularly Drs H.

Anderson and H.F. Glen, as well as Miss M. Koekemoer

for collecting specimens.

REFERENCES

ADANSON, M. 1763. Families des plantes 2. Paris.

ARNELL, S. 1956. Hepaticae collected by O. Hedberg etal. on the East

African mountains. Arkiv fur Botanik 3: 517-562.

ARNELL, S. 1963. Hepaticae of South Africa. Swedish Natural Science

Council, Stockholm.

Bothalia 23,1 (1993)

57

ASAKAWA, Y., TOKUNAGA, N., TOYOTA, M., TAKEMOTO, T,

& SUIRE, C. 1979. Chemosystematics of bryophytes I. The

distribution of terpenoids in bryophytes. Journal of the Hatton

Botanical Laboratory 45 : 395 —407.

ASAKAWA, Y., MATSUDA, R. & TAKEMOTO, T. 1980a. Mono- and

sesquiterpenoids from Wiesnerella denudata. Phytochemistry 19'

567-569.

ASAKAWA, Y., TOKUNAGA, N., TAKEMOTO, F., HATTORI, S.,

MIZUTAN1, M. & SUIRE, C. 1980b. Chemosystematics of bryo-

phytes IV. The distribution of terpenoids and aromatic compounds

in Hepaticae and Anthocerotae. Journal of the Hattori Botani-

cal Laboratory 47: 153 — 164.

ASAKAWA, Y., MATSUDA, R. & TAKEDA, R. 1981. Mono- and

sesquiterpenoids of Conocephalum supra decompositum.

Phytochemistry 20: 1423, 1424.

BENSON-EVANS, K. & HUGHES, J.G. 1954. The physiology of sexual

reproduction in Lunularia cruciata (L.) Dum. Transactions of

the British Bryological Society 2: 513— 521.

BERRIE, G.K. 1960. The chromosome numbers of liverworts (Hepaticae

and Anthocerotae). Transactions of the British Bryological Society

3: 688-705.

BEST, E.B. 1990. The Bryophyta of Zimbabwe — an annotated check-

list. Kirkia 13: 293-318.

BISCHLER, H. 1988. Relationships in the order Marchantiales

(Hepaticae). Journal of the Hattori Botanical Laboratory 64:

47-57.

BIZOT, M. & POCS, T. 1979. East African bryophytes. III. Acta Botan-

ica Academiae Scientiarum Hungaricae 25: 223—261.

BIZOT, M., POCS, T. & SHARP, A.J. 1985. Results of a bryogeographi-

cal expedition to East Africa in 1968, III. The Bryologist 88:

135-142.

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

upon cytogenetical and morphological evidence. Nova Hedwigia

45: 41-52.

BOWDICH, S. 1825. Excursions in Madeira and Pono Santo: 35.

CAMPBELL, E.O. 1965. Lunularia in New Zealand. Tuatara 13: 31—42.

CAMPBELL, E.O., MARKHAM, K.R., MOORE, N.A., PORTER,

L.J. & WALLACE, J.W. 1979. Taxonomic and phylogenetic

implications of comparative flavonoid chemistry of species in

the family Marchantiaceae. Journal of the Hattori Botanical

Laboratory 45: 185—199.

ENGEL, J.J. & SCHUSTER, R.M. 1982. Austral Hepaticae XV.

Brevianthaceae : a monotypic family endemic to Tasmania. The

Bryologist 85 : 375 — 388.

EVANS, A.W. 1919. A taxonomic study of Dumortiera. Bulletin of the

Torrey Botanical Club 46: 167—182.

GOODMAN, G.T. 1956. Sexual Lunularia cruciata (L.) Dum. in South

Wales. Transactions of the British Bryological Society 3: 98-102.

GOTTSCHE, C.M., LINDENBERG, J.B.G & NEES AB ESENBECK,

C.G. 1844—1847. Synopsis Hepaticarum. Hamburg.

GRIFFITH, W. 1849. On the higher cryptogamous plants. Notulae ad

plantas asiaticas, part 2: 285—352. Calcutta.

GROLLE, R. 1976. Verzeichnis der Lebermoose Europas und benach-

barter Gebiete. Feddes Repertorium 87: 171-279.

GROLLE, R. 1983. Nomina generica Hepaticarum, references, types

and synonymies. Acta Botanica Fennica 121: 1—62.

GROLLE, R. & PIIPPO, S. 1984. Annotated catalogue of western

Melanesian bryophytes 1. Acta Botanica Fennica 125: 1—86.

HASSEL DE MENENDEZ, G.G. 1963. Estudio de las Anthocerotales

y Marchantiales de la Argentina. Opera Lilloana 7: 1—297.

HEITZ, E. 1927. Uber multiple und aberrante Chromosomenzahlen. HI.

Das Genom der Lebermoose. Abhandlungen der naturwissen-

schaftlichen Verein zu Hamburg 21: 48—58.

HILL, J. 1773. A general natural history 2, edn 2: 120.

HOOKER, J.D. & TAYLOR, T. 1844. Flora antarctica. London Journal

of Botany 3 : 576.

HOWE, M.A. 1899. The Hepaticae and Anthocerotes of California.

Memoirs of the Torrey Botanical Club 7: 13—33.

INOUE, H. 1976. Illustrations of Japanese Hepaticae 2. Tokyo.

KLINGGRAEFF, H.E.M. 1858. Die hdheren Ctryptogamen Preussens.

Konigsberg.

KUNTZE, O. 1891. Revisio generum plantarum 2: 837.

LINDBERG, S.O. 1868. Musci novi scandinavici. Notiser Sdllskap pro

Fauna et Flora Fennica Fdrhandlingar 9 : 298.

LORBEER, G. 1934. Die Zytologie der Lebermoose mit besonderer

Beriicksichtigung allgemeiner Chromosomenffagen. Jahrbuch fur

wissenschaftliche Botanik 80: 567-818.

MACKAY, J.T. 1836. Flora hibemica. Dublin.

MACVICAR, S.M. 1926. The student’s handbook of British hepatics:

464. Eastbourne.

MICHELI, P.A. 1729. Nova plantarum genera. Florintiae.

MULLER, K. 1951—1958. Die Lebermoose Europas, in Rabenhorst’s

Kryptogamenflora 6: 416—471.

NECKER, N.J. DE 1790. Elementa: 345.

NEES AB ESENBECK, C.G.D. 1824. Dumortiera . In Reinwardt, Blume

& Nees ab Esenbeck, Hepaticae javanicae. Nova Acta Academiae

Caesareae Leopoldina-Carolinae Germanicae Naturae Curio-

sorum XII: 410.

NEES AB ESENBECK, C.G.D. 1838. Naturgeschichte der europdischen

Lebermoose 4. Berlin & Breslau.

NELSON, E.C. & PARNELL, J. 1992. Flora hibemica (1836): its

publication and aftermath as viewed by Dr Thomas Taylor. Taxon

41: 35-42.

PORTER, L.J. 1981. Geographic races of Conocephalum (Marchantiales)

as defined by flavonoid chemistry. Taxon 30: 739 -748.

RADDI, G. 1818. Novarum vel rariorum ex Cryptogamia stirpium in

agro Florentino collectarum Decades duae. Opuscoli scientifici

di Bologna 2: 349—361.

SAXTON, W.T. 1931. The life history of Lunularia cruciata (L.) Dum.,

with special reference to the archegoniophore and sporophyte.

Transactions of the Royal Society of South Africa 19: 259—268.

SCHIFFNER, V.F. 1893. Hepaticae bearbeitet von Dr Victor Schiffner,

aus Engler-Prantl. Die naturlichen Pflanzenfamilien 1: 3—141.

SCHUSTER, R.M. 1984a. Diagnoses of some new taxa of Hepaticae.

Phytologia 56: 65—74.

SCHUSTER, R.M. 1984b. New manual of bryology 2: 892-1070. The

Hattori Botanical Laboratory, Nichinan, Miyazaki, Japan.

SCHUSTER, R.M. 1992. The Hepaticae and Anthocerotae of North

America. Vol. 6.

SIM, T.R. 1926. The Bryophyta of South Africa. Transactions of the

Royal Society of South Africa 15: 1-475.

SMITH, j.E. 1833. Cryptogamia. The English Flora 5: 106.

SPRUCE, R. 1855. Hepaticae amazonicae et andinae. Transactions of

the Botanical Society, Edinburgh 15: 1-590.

STEPHANI, F. 1899. Species hepaticarum. Bulletin de VHerbier Boissier

7: 198-225.

SWARTZ, O. 1788. Nova genera et species plantarum: 145.

TATUNO, S. 1938. Uber Polyploidie und geographische Verbreitung bei

Dumortiera hirsuta. Botanical Magazine Tokyo 52: 434—441.

TATUNO, S. 1939. Weitere Untersuch ungen uber die Polyploidie und

die geographische Verbreitung bei Dumortiera hirsuta. 1—11.

Botanical Magazine Tokyo 53: 345—350.

TATUNO, S. 1941. Zytologische Untersuchungen liber die Lebermoose

von Japan. Journal Scientifica Hiroshima University 4: 73—187.

TAYLOR, T. 1836. De Marchantieis. Transactions oftheLinnean Society

17: 375 - 395.

VANA, J., POCS, T. & DE SLOOVER, J.L. 1979. Hepatiques

dAfrique tropicale. Lejeunia 98: 1—23.

VANDEN BERGHEN, C. 1965. Hepatiques recoltees par le Dr J.-J.

Symoens dans la region peri-Tanganyikaise. Bulletin de la Societe

Royale de botanique de Belgique 98: 129—174.

WHITTEMORE, A.T. 1991. The secondary chemistry of the Marchan-

tiales. Advances in Bryology 4.

WILLDENOW, C.L. 1809. Gesellschaft naturforschender Freunde,

Berlin. Magazin fur die neuesten Entdeckungen 3: 101.

.

.

'

Bothalia 23,1: 59-64 (1993)

Panicum simulans (Paniceae, Poaceae), a new species from southern

Africa and its leaf anatomy

L. SMOOK* and R.P. ELLIS**

Keywords: leaf anatomy, morphology, Namibia, new species, Panicum, Poaceae

ABSTRACT

Panicum simulans Smook from northern Namibia is formally described. The morphology and leaf blade anatomy in

transection and surface view, as observed under the light microscope, are described. The species is compared to others

with which it has been confused in the past, P. simulans can be readily distinguished from P. novemnerve Stapf and

P. schinzii Hack, on the basis of both anatomical and morphological characters such as the venation of the lower glume

and the photosynthetic anatomy, which is of the NAD-me subtype of the C4 photosynthetic pathway. On this basis it appears

that P. simulans is closely allied to the P. coloratura L. group of species and should be placed in section Panicum of subgenus

Panicum.

UITTREKSEL

Panicum simulans Smook vanaf die noordelike Namibie word formeel beskryf. Die morfologie en blaaranatomie, in deursnee-

en oppervlakaansig, soos met die ligmikroskoop waargeneem, word beskryf. Die morfologie en blaaranatomie van P. simulans

word vergelyk met die van twee ander spesies wat verwarring in die verlede geskep het. P. simulans kan maklik van

P novemnerve Stapf en P. schinzii Hack, onderskei word op grond van die bearing van die onderste gluma en die fotosintetiese

anatomie wat van die NAD-me subtipe van die C4-fotosintetiese weg is. Op grond van hierdie gegewens word P. simulans

in die seksie Panicum van subgenus Panicum geplaas.

Panicum simulans Smook, sp. nov.

Panicum sp. 2 (Giess 8605 ) in Gibbs Russell et al 242 (1990).

Panicum spp. in Group 3a in Ellis (1988).

Panicum simulans Smook, sp. nov., P. novemnervi Stapf

similis sed gluma inferiori uninervi, flosculo inferiori

masculo, paleaque inferiori bene evoluta differt; P. schinzii

Hack, similis sed nodis culmorum adpresso-hirsutis,

apicibus glumae superioris et lemmae inferioris erectis vel

recurvis, atque chloroplastis centripetis, non centrifugis,

differt.

Annual. Plant variable; usually yellowish green in

colour, often flushed purple. Culms erect to geniculate,

600(— 1 200) mm high (from plant base to top of inflores-

cence); sometimes rooting at nodes; nodes with adpressed

silvery white hairs pointing upwards; base covered with

leaf sheaths; internodes usually not visible. Leaf sheaths

glabrous or with bulbous-based hairs varying in length and

size of hair and relative size of base. Ligule a fringed

membrane 1.4 mm long (membranous rim 0.3 mm, cilia

1.1 mm long). Leaf blade linear, 5 — 200(— 250) x 2—15

mm, straight or cordate at base, flat, many-nerved,

bulbous-based hairs present or absent; apex acute to

acuminate; margins white, smooth to scaberulous to

densely scabrid; adaxial surface densely papillate

especially on lower leaves; nerves of abaxial leaf surface

smooth, occasionally densely scabrid especially near leaf

apex (variable on same plant).

* National Botanical Institute, Private Bag X101, Pretoria 0001.

** Agricultural Research Council, Roodeplaat Grassland Institute, Private

Bag X05, Lynne East, Pretoria 0039.

MS. received: 1992-02-13.

Inflorescence a panicle (80-)160-250(— 400) mm long,

narrowly obovate to obovate, occasionally oblanceolate,

sometimes asymmetrical (lowest branch longest,

protruding beyond other branches), moderately branched,

usually ascending and spreading; lowest branches single

or a number arising from same side, not whorled; next

set of branches often whorled or in a pseudo-whorl;

branches naked for a long way up from base with spike-

lets crowded at apices, smaller branches usually scabrid

with prickles becoming denser, longer and larger towards

spikelets; spikelets usually in pairs on long-short pedicels

on branchlets. Spikelets narrowly obovate to obovate-

elliptic, (2.2— )2.4-2.6(-3.0) mm long, acute to acumi-

nate, often flushed purple. Lower glume broadly ovate, '/3

to just over '/2 as long as intact spikelet, clasping base of

spikelet, membranous; central nerve distinct, often

scaberulous to scabrid; lateral nerves obscure, if present,

only visible at base; apex acute to acuminate, sometimes

minutely mucronate or with a brush of prickles. Rachilla

pronounced between glumes. Upper glume 2.4— 2.6(— 3.0)

mm long, longer than lower lemma, membranous, 7—9(11)-

nerved (nerves nearest margins sometimes obscure and

short); central nerve thickened at apex, forming a usually

scabrid mucro; apex acute to acuminate, either erect or

slightly recurved outwards; margins below apex mem-

branous and folding inwards, often flushed purple and then

mainly at apex. Lower floret male only; anthers 3, deve-

loped 1.4— 1.7 mm long or rudimentary, both conditions

present on same inflorescence, dark orange-brown. Low-

er lemma similar to, but shorter than upper glume, fitting

into its apex, (7-)9-nerved; cross venation often visible;

central nerve scaberulous, thickened towards apex; apex

acute to truncate; margins broad and membranous,

either folded in or out, often purple especially near apex.

Lower palea well developed, nearly as long and as wide

as lower lemma. Upper lemma and palea smooth, shiny,

pale, white to straw-coloured, occasionally dark with

60

Bothalia 23,1 (1993)

FIGURE 1. — Panicum simulans. A,

habit, x 0.5; B, node showing

adpressed hairs, x 2.5; C,

ligule, x 2.5; D, spikelet

showing lower glume, X 10;

E, spikelet showing upper

glume, X 10; F, lateral view of

open spikelet, x 10. From

Smook 5108 and Giess 8605.

distinct yellow nerves on upper lemma, both colour forms

sometimes present on same plant. Styles .2, separate;

stigmas plumose, purple to dark brown (on herbarium

specimens). Figure 1.

three species clearly and to justify the recognition of

P. simulans (Table 1). The specific epithet refers to the

superficial similarity between P. simulans, P. schinzii and

P. novemnerve.

TYPE. — Namibia, 1915 (Otjihorongo): 25 km N of

Outjo on road to Okaukuejo, (— DD), Smook 5108 (PRE,

holo.) ( Smook 5108a , PRE, isotype*).

Panicum simulans has been confused with P. schinzii

Hack, and P. novemnerve Stapf in the past, and the

specimens now included in this new species have been

misidentified as either of these two species. However, this

study shows that several characters serve to separate these

* This is the anatomical voucher specimen.

Figure 2 shows the known distribution of P simulans.

It is restricted to northern Namibia but may possibly also

occur in southern Angola. It is associated with various

vegetation types ranging from mopane woodland to

mopane mixed with Acacia, Combretum or Terminalia to

sweet grassland on limestone soils such as occur around

Etosha Pan.

Panicum simulans is hydrophytic and grows typically

in areas with seasonally high moisture regimes as found

in vleis and around fountains. It may even occur in more

Bothalia 23,1 (1993)

61

TABLE 1. Comparison between Panicum novemnerve, P. simulans and

P. schinzii

ephemerally moist situations such as in depressions in

disturbed situations along roadsides or irrigated lands or

in natural hollows in granite outcrops or saturated soil

pockets overlying calcrete.

Specimens examined

NAMIBIA. — 170 (Swartbooisdrift): Kaokoveld, (— DD), Merxmueller

& Giess 35538. 180 (Ohopoho): Kaokoveld, (— BB), De Winter & Leist-

ner 5160. 1815 (Okahakana): Etosha National Game Park, (-DC), Le

Roux 1399. 1817 (Tsintsabis): 56 km S of Namutoni on road to Tsumeb,

(—CD), Smook 5153*'. Farm Falkenhain GR/TS 303, (—DA), Giess 15090.

1914 (Kamanjab): 15 km N of Otjovasandu, (— AB), Giess 9253; Farm

Grootberg, (—CD), Du Toil 236. 1915 (Okaukuejo): Etosha National Game

Park, Vogelnestvley, (-BB), Giess & Mueller 13971 , 10 km S of

Okaukuejo, Giess & Loutit 14143; 9 km E of Okaukuejo, Smook 5109*.

(— BB), Ttnley 1299A; Ombika, (— BD), Le Roux 328; Ombika detour,

Giess & Loutit 14113; near Ombika, Giess 15098; 25 km N of Outjo on

road to Okaukuejo, (— DD), Smook 5108*. 1916 (Gobaub): Etosha National

Game Park, Olifantsbad, (— AA), Ellis 5271* 5273*; Gemsbokvlakte,

(— AA), Smook 5110*; Halali, (-BA), Van der Westhuizen 23. 1917

(Tsumeb): Tsumeb, (—BA), Giess 8605. 1918 (Grootfontein): Farm Sus,

(-AB), Schweickerdt 2144. 2015 (Otjihorongo): Farm Pamela, (-AB),

Volk 2855; Farm Babatsi, (— AB), Gibbs Russell & Smook 5628*; Farm

Straussenheim OUT 133, (— BB), Giess, Volk & Bleissner 6004. 2115

(Karibib): Farm Etemba, (-BC), Giess 10799; Farm Schlucht OM 162,

(-DA), Giess 8435; Farm Ameib, Giess, Volk & Bleissner 5898. De

Winter & Hardy 8072.

LEAF ANATOMY OF PANICUM SIMULANS

In order to obtain further evidence relating to the taxo-

nomic status and position of P. simulans, a detailed

anatomical study was undertaken. The methods used are

described in Ellis (1988) and the terminology used is

defined in Ellis (1976 & 1979). The following abbrevia-

tions will be used:

vb/s = vascular bundle(s)

l'vb/s = first-order vascular bundle(s)

2’vb/s = second-order vascular bundle(s)

3’vb/s = third-order vascular bundle(s)

ibs = inner or mestome sheath

obs = outer or parenchyma sheath

Leaf blade in transverse section

Outline : expanded blade, flat to very broadly V-shaped;

arms of lamina straight or gently undulating; blade

symmetrical about midrib. Ribs and furrows : shallow and

wide adaxial furrows present between all vbs; slight,

* voucher specimens for anatomical study

rounded ribs occur in association with all vbs; similar ribs

occur over all vbs. No abaxial ribs or furrows present.

Median vascular bundle; variable, either median bundle

only present (this indistinguishable structurally from lateral

l’vbs and without additional parenchyma tissue) (Figure

3A) or keel with associated colourless parenchyma tissue

present (Figure 3B); 1, 3 or 5 vbs incorporated in keel;

all bundles abaxially located; larger keels rounded with

single adaxial groove; no adaxial sclerenchyma developed

and abaxial girders associated with all vbs; no air spaces

in keel. Vascular bundle arrangement; 5, 7, 9 or 11 l’vbs

in transection; 3, 4 or 5 3’vbs between consecutive l’vbs;

narrower leaves without keels, with 5 l’vbs and with three

3’vbs between successive larger bundles. No 2’vbs. All

vbs situated in centre of blade. Vascular bundle descrip-

tion; 3’vbs circular in outline with xylem and phloem

distinguishable, l’vbs circular in shape, lysigenous cavities

present (Figure 3E, F) and metaxylem vessels narrow

(diameters less than those of obs cells) and circular.

Vascular bundle sheaths; obs conspicuous, round, entire

(except with small abaxial interruption in l’vbs) and

without extensions; consists of 6, 7 or 8 cells around 3’vbs

and 11—14 cells around l’vbs; obs cells fan-shaped with

radial walls straight and outer tangential walls inflated

(Figure 3E, F); all cells similar in size and shape;

chloroplasts dense, filling entire cell lumen or concentrated

centripetally, near inner tangential wall. Ibs absent in

3’vbs; complete but inconspicuous in l’vbs; cells very

much smaller than obs cells; adaxial cells slightly larger

and thin-walled; lateral and abaxial cells uniformly

thickened but degree of secondary thickening very vari-

able. Sclerenchyma; minute adaxial and abaxial strands

only associated with l’vbs; no fibrous tissue in associa-

tion with 3’vbs. Small sclerenchyma caps in leaf margin

(Figure 3A). Chlorenchyma; distinctly radiate; single layer

of tabular cells completely surrounds all vbs (except some-

times for a short abaxial interruption with l’vbs); radiating

chlorenchyma of successive bundles in direct contact and

not separated by colourless parenchyma. No colourless

parenchyma tissue present (except in midrib, if present).

Adaxial epidermal cells; bulliform cells small, in rather

extensive groups; central bulliform cell occupies less than

'/4 of leaf thickness; cuticle very thin; no macrohairs,

except rarely in association with margin; no prickles or

hooks evident; cuticular papillae common, much narrower

than epidermal cells and with one per cell as seen in

FIGURE 2. — The known distribution of Panicum simulans.

62

Bothalia 23,1 (1993)

FIGURE 3. — Leaf blade anatomy of Panicum simulans as seen in transverse section. A, outline of specimen without keel; median vascular bundle

only (arrowed), this bundle structurally indistinguishable from other first-order vascular bundles; B, outline of specimen with distinct keel

incorporating four vascular bundles and adaxial colourless parenchyma tissue; C, typical C4 photosynthetic anatomy with prominent paren-

chyma sheaths; D, specimen showing distinct centripetal location of Kranz chloroplasts in outer bundle sheath cells; E, dense chloroplasts

in outer bundle sheath cells and radiate chlorenchyma, note adaxial papillae; F, typical NAD-me type photosynthetic anatomy. A, B, x

100; C, D, X 250; E, F, x 400. A, C, Smook 5108', B, Smook 5720; D, Gibbs Russell & Smook 5268', E, Smook 5153', F, Smook 5110.

section (Figure 3C— F). Abaxial epidermal cells: central

cells of intercostal zones larger and resemble bulliform

cells in transection; cuticle thin; no macrohairs, prickles

or papillae.

Abaxial epidermis in surface view

Zonation: evident but costal zones overlying 3’vbs often

without silica bodies as a result of absence of sclerenchyma

strands in association with these bundles. Intercostal long

cells: elongated (length more than 3 X longer than width),

side walls parallel to outwardly bowed, end walls vertical;

anticlinal walls very thin and slightly undulating, with

degree of sinuosity varying with plane of focus; cell shape

varies across individual intercostal zones with two files

of bulliform- like cells per intercostal zone; these bulliform-

like cells actually represent central files of two intercostal

zones but costal zones are poorly differentiated (Figure

4C, D). Very few tall and narrow short cells present.

Stomata: 2 or 4 files of low dome-shaped stomata per

intercostal zone; usually separated by a single elongated

interstomatal cell. Papillae: absent. Prickles: absent.

Microhairs: bicellular with dehiscent distal cell; only basal

cells remain, basal cell elongated with length much greater

than width; occur in central files of intercostal zones

(Figure 4D). Macrohairs: absent. Silica bodies: very

irregular dumbbell-shaped; horizontally elongated; same

width as adjacent long cells. Costal short cells : 1 or 3 files

of cells comprise costal zones; central file with silica bod-

ies, these alternate irregularly with short to long short

cells. Costal zones often not associated with 3’vbs, only

present over l’vbs.

DISCUSSION

In general appearance P simulans closely resembles

P. schinzii , particularly those forms of both taxa that have

rounded or cordate leaf bases. However, the hairy nodes

(Figure IB) and pointed apex of the spikelets (Figure ID,

E) and colour of plant (yellowish green vs. green) of P.

simulans clearly separate these two taxa (Table 1) (Gibbs

Russell et al. 1990).

Bothalia 23,1 (1993)

63

All specimens from Namibia previously assigned to

P. schinzii have now been re-identified as P. simulans.

There is one exception, Volk 2672. However, there is no

habitat information for this specimen, and it may have been

a weed from a cultivated land and imported with the crop.

Panicum schinzii and P simulans both tend to be hydro-

phytic, but they are clearly separated geographically. Their

natural populations were undoubtedly originally allopatric,

but it can be expected that, with increasing agriculture

under irrigation, P. schinzii will become established in the

range of P. simulans.

In addition to the morphological differences between

Panicum simulans and P. schinzii, there are major differ-

ences in the photosynthetic leaf anatomy between these

two taxa. P. simulans has the NAD-me subtype of the C4

photosynthetic pathway, as the anatomy is Kranz with a

double bundle sheath with the specialized chloroplasts of

the parenchyma sheath centripetally arranged (Ellis 1988).

The Panicum species with this type of photosynthetic

anatomy constitute the ‘true’ Panicum species (Brown

1977) as P. miliaceum L., the type of the genus, belongs

to this group.

Panicum schinzii, on the other hand, characteristi-

cally has the Kranz chloroplasts located against the outer

tangential cell wall. These chloroplasts are relatively small

and often form only a thin peripheral layer in contrast to

the large, dense chloroplasts of P. simulans. The anatomy

of P. schinzii is typical of that of the PEP-ck subtype of

the C4 pathway (Ellis 1988). Brown (1977) considered all

the Panicum species of this PEP-ck subtype, as doubtful

members of the genus and as belonging to Brachiaria,

together with Urochloa and Eriochloa. This applies

particularly to those species with rugose lemmas.

Ellis (1988) grouped Panicum simulans and P. schinzii

in two different subgeneric groups because of these distinct

anatomical differences. P. schinzii was placed in Group

2d whereas P. simulans (as Panicum spp., together with

P. novemnerve) was placed in Group 3b.

Group 2, although the leaf anatomy is typically PEP-ck

throughout, actually consists of two distinct and separate

groups. Group 2d with P. schinzii, together with P. gilvum

Launert, P. impeditum Launert, P. repens L. and P. sub-

albidum Kunth, are all hydrophytic and have pale, glossy

upper lemmas. These taxa are clearly not closely allied

to Groups 2a— 2c, all of which appear to belong to the

panicoid grasses with rugose lemmas. The P. schinzii

group of species (except P. repens) belongs to section

Dichotomiflora (Hitchc. & Chase) Honda of subgenus

Panicum (Zuloaga 1987) and is not closely allied to

P. simulans despite the superficial morphological resem-

blance.

The work of Ohusigi et al. (1982) urges caution in

assigning P. schinzii and its allies to the PEP-ck photosyn-

thetic subtype as they may actually all be NAD-me. The

biochemical typification of these taxa urgently needs

verification before final taxonomic decisions can be made.

However, this has no bearing on the separation of P.

simulans from this group.

In the past P. novemnerve has also been confused with

P. simulans. However, P. simulans has only one distinct

FIGURE 4.— Abaxial epidermal structure of Panicum simulans. A, typical zonation pattern with 5 narrow costal zones located over third-order

bundles between successive first-order bundles; B, leaf margin showing cushion-based macrohair; C. stomata and microhairs in intercostal

zones; D, low dome-shaped stomatal subsidiary cells, basal cells of microhairs and very irregularly dumbbell-shaped silica bodies. A,

B, x 160; C, x 250; D, x 400. A, D, Smook 5153 ; B, Smook 5U0; C, Smook 5109.

64

Bothalia 23,1 (1993)

and obvious nerve on the lower glume (Figure ID) instead

of 3—5 as in P novemnerve. In P. novemnerve the lower

floret is sterile with a reduced palea (Stapf 1920), whereas

in P simulans the lower floret is male with a well-

developed palea (Gibbs Russell et al. 1990). These two

taxa can, therefore, be readily separated on morphological

criteria.

Panicum simulans and P. novemnerve also share similar

damp habitats although P. novemnerve appears to prefer

shady areas. They are sympatric in Namibia, but P. novem-

nerve also occurs in Botswana and the Transvaal.

The leaf anatomy of P novemnerve (as based on Smook

5163 and Gibbs Russell & Smook 5235) and P. simulans

is virtually identical. Both have the NAD-me type anatomy

with very little sclerenchyma development. The only

discernible difference appears to be that the intercostal long

cells in P. novemnerve are often separated by tall and

narrow short cork cells, which is very seldom the case

in P. simulans. Both P simulans and P. novemnerve

specimens were included under Panicum spp. of Group

3a by Ellis (1988) because of this similarity in leaf blade

anatomy. These two annual species, together with P.

arcurameum Stapf and P. atrosanguineum A. Rich.,

undoubtedly belong to the P. coloratum L. group of

species, and show very close similarities with P. pil-

gerianum (Schweick.) Clayton [= Psilochloa pilgeriana

(Schweick.) Launert], another annual hydrophyte. This

latter species has only recently been placed in Panicum ,

and this small group of annual species appears to form

a clear subgrouping within the P coloratum group of

species. They resemble an unnamed perennial entity within

P. coloratum particularly closely, as discussed by Ellis

(1988). Specimens of this taxon are Ellis 1783, 2905 and

2912, all of which are also hydrophytes. Oryzidium

bamardii C.E. Hubb. & Schweick., may also belong to

this assemblage.

Panicum simulans, therefore, appears to belong with the

‘true’ Panicum species of section Panicum of subgenus

Panicum (Zuloaga 1987). These are typical NAD-me

species as suggested by the leaf anatomy but biochemical

typing is required.

REFERENCES

BROWN, W.V. 1977. The Kranz syndrome and its subtypes in grass

systematics. Memoirs of the Torrey Botanical Club 23: 1—97.

ELLIS, R.P. 1976. A procedure for standardizing comparative leaf blade

anatomy in the Poaceae. I. The leaf blade as viewed in trans-

verse section. Bothalia 12 : 65-109.

ELLIS, R.P. 1979. A procedure for standardizing comparative leaf blade

anatomy in the Poaceae. II. The epidermis as seen in surface view.

Bothalia 12: 641—671.

ELLIS, R.P. 1988. Leaf anatomy and systematics of Panicum (Poaceae:

Panicoideae) in southern Africa. Monographs in Systematic

Botany IS: 129-156.

GIBBS RUSSELL, G.E., WATSON, L., KOEKEMOER, M., SMOOK,

L., BARKER, N.P., ANDERSON, H.M. & DALLWITZ, M.J.

1990. Grasses of southern Africa. Memoirs of the Botanical Survey

of South Africa No. 58.

OHUSGI, R., MURATA, T. & CHONAN, N. 1982. C4 syndrome of

species in the Dichotomiflora group of the genus Panicum.

Botanical Magazine (Tokyo) 95: 339—347.

STAPF, O. 1920. Gramineae. In D. Prain, Flora of tropical Africa, Vol.

9. Reeve, Ashford.

ZULOAGA, F.O. 1987. Systematics of New World species of Panicum

(Poaceae: Paniceae). InT.R. Soderstrom et al., Grass systematics

and evolution: 287—306. Smithsonian Institution Press, Washing-

ton D.C.

Bothalia 23,1: 65-83 (1993)

Notes on African plants

VARIOUS AUTHORS

ROSACEAE

OBSERVATIONS ON CLIFFORTIA MICRANTHA

A fresh specimen of a Cliffortia species from the Swart-

berg ( Vlok 2113) clearly showed more than one style and,

following Weimarck’s (1934, 1948) keys, was identified as

C. propinqua Eckl. & Zeyh. (subgenus Digraphidium sec-

tion Complanatae) , but its appearance differed greatly

from that species. It did, however, match C. micrantha

Weim. (subgenus Monographidium, section Costatae or

Bacciformes ; see below) a single-styled species, both in

appearance and by key except for the number of styles.

With the handling of the fresh specimen, it was clear that

the styles were easily shed. A re-evaluation of existing

herbarium collections of C. micrantha seemed, therefore,

necessary. In 15 herbarium specimens examined from three

major Cape herbaria (BOL, NBG, STE), two-styled fruits

were found occasionally on only five of the specimens;

most fruits had no styles and a few only one. The fresh

specimen ( Vlok 2113) was predominantly two-styled with

only a few three-styled fruits.

According to Weimarck (1934, 1948), the number of

styles and achenes correspond. It should therefore be

feasible to count the achenes instead of the styles.

The number of achenes in at least two fruits per

specimen, were counted for the 15 specimens of C.

micrantha. The fruits of the fresh Swartberg specimen

( Vlok 2113) were also examined for number of achenes.

Esterhuysen 28523, identified by the collector as C.

micrantha, but possibly of hybrid origin (see below), was

also examined.

All the fruits of C. micrantha proper contained at least

two achenes. A single specimen from near Prince Albert

{Bond 1680) had two to three achenes per fruit. A

specimen from Noukloof Nature Reserve {Laidler 154) had

two to four achenes per fruit as well as two styles on some

of the fruits. The Swartberg specimen ( Vlok 2113) had two

to three achenes corresponding with the number of styles

(Figure LA— E). The fruit of Esterhuysen 28523 appeared

more oblong and furrowed than the norm for C. micran-

tha and contained only one achene. Closer examination

showed this specimen to differ from C. micrantha in the

shape and size of the leaves as well, tending towards the

characters found in C. cervicomu Weim. (Weimarck 1959).

If it is accepted that one style concurs with one achene

and two (or more) styles with two (or more) achenes con-

sistently enough to subdivide a genus (Weimarck 1934,

1948), then C. micrantha must be regarded as a multiple-

styled species because there were never less than two

achenes per fruit in all the specimens examined. This is

confirmed by the fresh Swartberg specimen {Vlok 2113)

which has two to three styles and two to three achenes.

Weimarck’s (1934, 1948) perception of C. micrantha as

having a single style could be due to the fact that the styles

in his material had been shed. As he equated the number

of achenes to the number of styles, it is most likely that

he did not examine the achenes in this species.

Weimarck (1940) placed C. micrantha in the section

Costatae with C. serpyllifolia Cham. & Schlechtd. and

C. browniana Burtt Davy on the basis of its ribbed fruits

and the mistaken assumption that it has a single style.

C. micrantha can, however, have ribbed or smooth fruits,

depending on the age of the fruit, (Figure LA, F & G) and

was therefore placed in two divisions in his key for the

section Costatae (Weimarck 1948). In the same article,

he placed C. micrantha in his key for the section

Bacciformes as well (Weimarck 1948). He discussed it

under the latter section which contains only one other

species, C. baccans Harv. This latter species has a smooth

and berry-like fruit, resembling the mature fruit of

C. micrantha, but with only one style. If C. micrantha

is retained in either the sections Bacciformes or Costatae,

the number of styles loses its taxonomic importance.

Furthermore, the feasibility of subdividing the genus into

subgenera on this basis becomes questionable.

On the above evidence, C. micrantha should be placed

in the subgenus Digraphidium on the basis of the

predominance of two-styled flowers. This placement is fur-

ther substantiated by the trifoliate leaves, the tetra-

merous flowers and the low stamen count common in this

subgenus. However, fruits of the only section {Complana-

tae) of this subgenus are flattened and hard, and not

berry-like as are the mature fruits of C. micrantha (Figure

LA). The older fruits, however, can appear ribbed and the

two-achened ones are dorsiventrally flattened, to some

extent resembling the fruits in the section Complanatae

(Figure IF & G). C. micrantha might therefore be placed

in the subgenus Digraphidium but in its own section. This

needs further investigation.

The possible occurrence of more than two styles and

achenes in the recognised two-styled species of Cliffortia

also needs investigation.

The probability of the specimen Esterhuysen 28523

being a hybrid between C. micrantha, a two-styled species,

and C. cervicomu Weim., a one-styled species, raises

more questions about relationships. A field trip to collect

fresh material of this probable hybrid was unsuccessful,

but needs to be done again as part of further work on

hybridisation and relationships in the genus.

SPECIMENS EXAMINED

For number of styles

CAPE. — 3320 (Montagu): Touwsberg, in kloof, S slope near top, 1 100

m, 11-07-1941, (-DB), Levyns 7483 (NBG). 3321 (Ladismith): Swart-

FIGURE I. — Cliffortia micrantha : A, mature fruit with two styles (bracteoles in place, covering the stipe). B, mature fruit in longitudinal section

showing flat sides of two achenes (a, a’) covered by two layers of tissue; stipe (b) attached; sepals, styles and bracteoles removed. C, top

view of three-styled fruit; D, styles from C; E, achenes from C; F, old fruit, side and corresponding top views showing ribs and dorsiventral

flattening; G, old fruit, showing four major and four minor ribs in a regular pattern. A— E, Vlok 2113 (STE); F, Laidler 154 (STE); G,

drawn from Boshoff 315 (STE). Scale bar = 2 mm.

berg, top of pass into Gamkaskloof, 1 333 m, 8-05-1963, (— BC), Taylor

4744 (STE); Swartberg, in ravine between Kliphuisvlei and Gamkas-

kloof, 1 250 m, 7-05-1989, (-BD), Vlok 2113 (PRE, STE); Noukloof

Nature Reserve, gentle SE slope, 566 m, 12-07-1982, (— CA), Laidler

154 (STE); Roodeberg, S slope near stream, 24-05-1950, (— CB),

Esterhuysen 17152 (NBG).

For number of achenes

CAPE. — 3320 (Montagu): Anysberg, S slopes, 810 m, 2-08-1956,

(-DA), Wurts 1424 (NBG); Touwsberg, steep rocky lower S slopes to

1 000 m, 1-06-1956, (-DB), Esterhuysen 25933 (BOL); Touwsberg, 1 100

m, 11-07-1941, (-DB), Levyns 7483 (Type) (BOL, NBG, STE). 3321

(Ladismith): Ladismith, koppie 4 miles from town, 566 m, 15-08-1948,

(—AD), Levyns 9030 (BOL); Gamka Mtn Reserve, N slope (moist kloof),

866 m, 03-1976, (— BC), Boshoff P315 (STE); Prince Albert— Gamkaskloof

road, 890 m, 3-08-1979, (— BD), Bond 1680 (STE); Swartberg, in a ravine

between Kliphuisvlei and Gamkaskloof, 1 250 m, 7-05-1989, (— BD),

Vlok 2113 (STE); Noukloof Nat. Res., gentle SE slope, 566 m, 12-07-1982,

(-CA), Laidler 154 (STE); Roodeberg, S slopes near stream, 24-05-1950,

(— CB), Esterhuysen 17152 (BOL, NBG,); Roodeberg, 1 000 m,

10-08-1948, (-CB), Levyns 8981 (BOL, STE).

Probable hybrid (C. micrantha X C. cervicomu)

CAPE. — 3321 (Ladismith): Swartberg foothills between Bosluis Pass

and Die Hel, 10-10-1960, (-BC), Esterhuysen 28523 (BOL).

REFERENCES

WEIMARCK, H. 1934. Monograph of the genus Cliffortia. Gleerupska

Universitet Bokhandeln, Lund.

WEIMARCK, H. 1940. Cliffortia micrantha H. Weimarck spec. nova.

Botaniska Notiser 1940: 97—99.

WEIMARCK, H. 1948. The genus Cliffortia, a taxonomic survey.

Botaniska Notiser 1948: 175—185.

WEIMARCK, H. 1959. Four new Cliffortia species. Botaniska Notiser

112 : 77-79.

A.C. FELLINGHAM*

* Stellenbosch Herbarium, National Botanical Institute, P.O. Box 471,

Stellenbosch 7599.

MS. received: 1992-03-16.

Bothalia 23,1 (1993)

67

ROSACEAE

CLIFFORT1A FASCICULATA, A SUPERFLUOUS NAME FOR C. AMPLEX1STIPULA

In his monograph of the genus, Weimarck (1934) placed

Cliffortia amplexistipula Schltr. under the imperfectly

known species. He quoted Schlechter’s original descrip-

tion in full (Schlechter 1900) but pointed to the lack of

flower and fruit details which did not enable him to identify

the species accurately.

Cliffortia amplexistipula Schltr. in Botanische Jahr-

biicher 27: 140 (1900); Weim.: 157 (1934), sub species non

sat. cog. Type: In sabulosis prope Ezelbank, in montibus

Cederbergen, alt. c. 4000 ped., 2 Sept. 1896, Schlechter

8825 (Bf, holo. ; BOL! lectotype here designated; BM,

GRA, K, isolecto.).

This inadequacy led to the description of C. fasciculata

Weim. (Weimark 1946). Two years later, Weimarck (1948:

174) stated in one short sentence: ‘C. fasciculata has,

however, proved to be identical with C. amplexistipula

Schltr.’ and proceeded to use the latter name in his key.

This fact was also communicated to the Bolus

Herbarium as seen in a handwritten note on the sheet of

an isotype of C. fasciculata Weim. in BOL which reads:

‘Weimarck states in a report on material sent, that C.

fasciculata Weim. is a synonym for C. amplexistipula

Schltr.’ This note is initialled by E. Esterhuysen and dated

10/48.

In Gibbs Russell etal. (1987), both C. fasciculata Weim.

and C. amplexistipula Schltr. are listed as correct names.

Weimarck (1934) examined Schlechter’s type of C.

amplexistipula , ( Schlechter 8825), from five different

herbaria, the one in Berlin presumably being the holo-

type. As this one was destroyed, I hereby choose the Bolus

Herbarium specimen as lectotype of C. amplexistipula

Schltr.

C. fasciculata Weim.: 40 (1946). Type: N Cederberg, between Henning

(Heuning?) Vlei and Koupoort, 21-10-1945, Esterhuysen 12116 (LD, holo.;

BOL!, iso.).

Schlechter’s (1900) original description of C. amplexi-

stipula (as quoted by Weimarck 1934) did not include

flowers and fruit. Weimarck’s (1946) description of C.

fasciculata does include male and female flowers but no

fruit.

Description : to augment the description of this species,

the following observations are made:

Leaf sheaths: ventral apical portion with persistent red

fringe resembling stigma, amplexicaul when young, some-

times splitting when older. Stipules minute when young

to 0.25 mm long later. Leaflets: dorsal surface with raised

round spots to the margins (‘marginibus subscabridis’ of

Schlechter), spathulate with mucro minute and ventral in

young stage, to elliptic with mucro apical and well deve-

loped when mature (Figure 2A & B). Female sepals: 3,

1.0— 1.2 x 0.6— 0.7 mm, fleshy, broadly elliptic, acute,

mucronate, glabrous, dorsally with a few small round

raised spots (Figure 2C). Fruit: (2.6)2 .7 x 0.7(0.8) mm.

FIGURE 2. — Cliffortia amplexistipu-

la, Oliver 10046 (STE): A,

adaxial view of young leaf,

short shoot bearing young fruit

(C) removed; B, adaxial view

of older leaf, sheath opened; C,

young fruit with calyx, stigma

and bracts still attached; D,

mature fruit. Scale bars = 1

mm.

68

Bothalia 23,1 (1993)

narrowly oblong, slightly curved, 7(8)-ribbed with two ribs

flattened and extended around base forming con-

tinuous narrow wing, surface minutely scabrid to papillate

in places (Figure 2D).

Distribution and habitat : C. amplexistipula occurs on

the drier inland mountains, from the Kamiesberg in the

north to the Anysberg in the Little Karoo, in arid fynbos

on sandstones and quartzites, often on southern slopes or

in the lee of a rock.

Specimens examined in addition to the types

CAPE. — 3018 (Kamiesberg): Namaqualand, Welkom, Khamiesberg

near Garies, 1 100 m, 16-10-1954, (-AC), Esterhuysen 2J71I (BOL). 3118

(Vanrhynsdorp): Gifberg, 480 m, 14-07-1974, (—DC), Thompson 2088

(STE). 3119 (Calvinia): Calvinia Lokenburg, arid fynbos of

TMS ridges, 770 m, 28-08-1958, (— CA), Acocks 19719 (BOL); Lokenberg,

on rugged sandstone hills, 31-05-1964, (— CA), Esterhuysen 30707 (BOL).

3218 (Clanwilliam): South Cold Bokkeveld, Ceres, 4-10-1940, (— AB),

Bond 662 (NBG); Baviaansberg, Ceres, 1 450 m, 2-01-1942, (—BA),

Compton 12877 (NBG). 3219 (Wuppertal): Cederberg, Heuning Vlei,

29-12-1941, (-AA), Esterhuysen 7468 (BOL); Clanwilliam Div., Ceder-

berg, Tafelberg, shale band, 25-09-1942, (-AC),

Esterhuysen 8092 (BOL); Clanwilliam Div., foot of Tafelberg, shale band,

1 500 m, 29-12-1947, (-AC), Esterhuysen 14326 (BOL); Clanwilliam

Div., Duivelsgat, S Cederberg (Sneeuwberg area), 11-10-1946, (— CA),

Esterhuysen 13101 (BOL); Ceres Div., Zuurvlakte, E of Bokkeveld

Sneeuwkop area, 20-04-1946, (—CD), Esterhuysen 12744 (BOL, NBG,

SAM); Ceres, Stompiesfontein, Swartruggens, streambanks, 1-11-1961,

(—DC), Esterhuysen 29305 (BOL); Ceres, Stompiesfontein, Swartrug-

gens, rocky plateau, 1 100 m, 1-11-1961, (-DC), Esterhuysen 29327

(BOL). 3319 (Worcester): Gydo, Ceres, 1 000 m, 10-11-1946, (-AB),

Compton 18752 (NBG); Cold Bokkeveld, Winkelhaak area, flats NW

of Jakkalsdam, 960 m, 13-02-1992, (— AB), Oliver 10046 (STE). 3320

(Montagu): Laingsburg, Witteberg, on stony rocky south slopes, 1 300

m, 31-01-1961, (—AD), Esterhuysen 28867 (BOL, NBG); kloof on S slopes

of Anysberg, Little Karoo, 22-05-1950, (-DA), Esterhuysen 17077

(BOL); Anysberg, summit plateau, in lee of rock, 10-08-1991, (—DA),

Fellingham 1537 (STE).

REFERENCES

GIBBS RUSSELL, G.E. etal. 1987. List of species of southern African

plants, edn 2, part 2. Memoirs of the Botanical Survey of South

Africa No. 56: 64.

SCHLECHTER, R. 1900. Plantae Schlechterianae novae vel minus

cognitae describuntur. II. In A. Engler, Beitrage zur Flora von

Afrika. XVIII. Botanische Jahrbucher 27: 86—220.

WEIMARCK, H. 1934. Monograph of the genus Cliffortia. Gleerupska

Universitets Bokhandeln, Lund.

WEIMARCK, H. 1946. Further notes on the genus Cliffortia. Botaniska

Notiser 4 : 407—420.

WEIMARCK, H. 1948. The genus Cliffortia , a taxonomic survey.

Botaniska Notiser 2: 167—202.

A.C. FELLINGHAM*

* Stellenbosch Herbarium, National Botanical Institute, P.O. Box 471,

Stellenbosch 7599.

MS. received: 1992-03-16.

FABACEAE

VIGNA KOKH. A NEW SPECIES FROM SOUTHERN AFRICA

Vigna kokii B.J. Pienaar, sp. nov. (Sect. Micro-

spermae— Papilionoideae), V mudeniae B.J. Pienaar facie

caulibus pedunculis distincte costatis alatis similis;

V. richardsiae Verde. (Sect. Microspermae) stipulis reflexis

auriculis lateraliter libris caule connatis similis; a V

monophylla Taub. (Sect. Haydoniae) glandibus basalibus

verticillo interiore antherarum absentiis differt; styli

extensio ‘tumore’ in textura reducta ubi versus stigma

laterale flexa (non ut in V mudenia extensione ‘digito

primo’ et in V. monophylla tota absentia); reticulum extinae

granorum pollinis vix manifestum.

TYPE. — Transvaal, 2530 (Lydenburg): Schagen, wood-

land 0.8 km from Crocodile Hotel on road to Rosehaugh,

Pienaar 1364 (PRE, holo.; K, P, iso.).

Rootstock carrot-shaped, at length woody. Stem erect in

youth, twining at length, somewhat ridged to alate, scarce-

ly strigose to patent with light hairs. Leaflets oblong with

apex and base rounded in juvenile stage, ± 25—58 x

0.8-30.0 mm, surface smooth to touch, adnately strigose,

thickest along dorsal nerves and lamina margin, rhom-

boid to sublobular with rounded lobes at maturity, ± 80

x 60—70 mm at broadest point, apices abruptly narrowed,

obtuse, apiculate, base cuneate to obtuse, papyraceous,

strigose. Stipules with cordate base, laterally auriculate

at maturity, lanceolate, reflexed, ± 6.0 X 1.5 mm,

acuminate, margins ciliate; in youth base more or less

elongated, truncate (cf. V. mudenia B.J. Pienaar 1991).

Inflorescence contracted, peduncles alate, bearing at

apex one pair of yellowish green flowers, flushed violet,

with vertical row of extrafloral nectary glands between

them (Figure 3B). Calyx campanulate, lobes, ± 2 mm,

acute with broad bases, ± as long as tube or shorter, tube

up to 3.25 mm long, upper pair of lobes connate for '/3

to their length, strigose. Standard ovate to oblate,

emarginate, ± 10—14 mm long, spur straight with inferior

callosities divergent, auricular, superior callosities diver-

gent, narrow (vestigial). Wings narrowly ovate, ± 10.0 x

5.5 mm, auricle slender with raised cell sculpturing

(almost papillate). Keel ± 11 mm long, obtuse, slightly

upturned. Filament tube ± 10 mm long, free filaments

in two whorls, 3 or 4 mm long, anterior filament genicu-

late at base, ± 13 mm long; anthers oblate, glands at

base absent. Ovary pubescent, ± 9-10 mm long, style

broadened and flattened as it curves upward, pollen brush

on anterior face, apically twisted at maturity, accentuated

protuberance of style prolongation absent but stylar tissue

swollen as it bends toward stigmatic papillae at somewhat

more than 90°. Pollen grains scarcely reticulate, muri low,

rounded. Legumes beaked, + 68 mm long, scabrid with

long, stiff hairs along margin, 18-seeded. Seeds 3.4 mm

long, yellowish brown to dark brown, black mottled around

hilum, hilum scarcely eccentric, aril scarcely developed

or absent. Figure 4.

NAMIBIA. — 1724 (Katima Mulilo): 20 km SW of Katima Mulilo,

(— CA), De Winter 9180 (PRE). 1821 (Andara): Caprivi side of river,

near Andara Mission Station, (— AB), De Winter & Marais 4812 (PRE);

Dico, Andara, (-AB), Giess 15572 (WIND).

Bothalia 23,1 (1993)

69

FIGURE 3. Vigna kokii, Pienaar 1364 (PRE): A, style prolongation, reduced to a swelling, x 33; B, extrafloral nectaries, x 30; C, pollen

grain, X 458.

TRANSVAAL. — 2431 (Acomhoek): Nwanetzi, Kruger National Park,

(—AD), Coetzee 6062 (PRE). 2527 (Rustenburg): mountainside,

Saulspoort, (— AA), Germishuizen 514 (PRE). 2530 (Lydenburg):

Schagen, Farm of J.J. van Niekerk, Nelspruit District, (— BD), Liebenberg

3297 (PRE); 0.8 km from Crocodile Hotel on turnoff to Rosehaugh,

(-BD), Kok & Pienaar 1297 ( PRE, PRU), Pienaar 1364 (PRE).

Found in dense undergrowth of dry woodland in the

Transvaal and in Caprivi in northeastern Namibia. Figure

5.

In general facies V. kokii is very similar to V. mudenia

B.J. Pienaar (1991), with the stems and peduncles distinctly

alate and the contracted inflorescence bearing two yellow-

green flowers, but the leaflets are oblong with rounded

base and apex in youth, becoming rhombic or sublobular

at maturity. V kokii is similar to V richardsiae Verde.

(Sect. Microspermae ) in having stipules reflexed, connate

with the stem but with auricles laterally free; it differs from

V monophylla Taub. (Sect. Haydonia) in the absence of

basal glands on the interior whorl of anthers; the style

prolongation (Figure 3A) is reduced to a ‘swelling’ in the

tissue of the style where it bends toward the lateral stigma,

as opposed to the ‘finger tip’ protuberance of V mudenia

and total absence of any prolongation in V monophylla-,

the exine sculpture of the pollen grains (Figure 3C),

appears more rounded than that of V. mudenia as opposed

to the total loss thereof in V monophylla . These charac-

ters support the decision to keep it in the same section

as V mudenia but to accept it as a new species. Its

distribution also differs from that of V mudenia.

The stylar thickening (rather than a true protuberance),

at the bend toward the lateral stigma, may represent an

evolutionary stage more advanced than in other members

of the section Microspermae. This trend probably reflects

an incipient stage toward the loss of the stylar beak, as

displayed in the subgenus Haydonia (Pienaar 1992). The

reticulation of the pollen grains has almost disappeared,

a further development toward the eventual smoothness of

the exine in section Haydonia (e.g. V. monophylla Taub.).

a^lPiewAPR.

FIGURE 4. — Vigna kokii. Pienaar 1344 (PRE): A, leaf, X 1; B, stipule, X 3; C, calyx, x 3; D, standard with two pairs callosities, X 4; E,

wing, x 4; F, anther, X 7; G, style prolongation (s), stigma (st), pollen brush with pollen (p), X 10; H, legumes on peduncle, X 1;

I, legumes, X 1; J, underground tuber, vertical, x 0.25.

70

Bothalia 23,1 (1993)

FIGURE 5. — Known distribution of Vigna kokii in southern Africa.

It gives me great pleasure to name this new species after

Prof. P.D.F. Kok, under whose guidance my taxonomic

work on the genus Vigna was initiated. One of the first

collections (in 1934) of the new species is Liebenberg 3297

(PRE), from a farm in the Schagen District. Prof. Kok

accompanied me on the first field trip to the Schagen area

in search of more material of the new species. He searched

as hard as I did until an immature plant was found in the

thickets and I was able to return to the locality a season

later. He also financed the trip.

ACKNOWLEDGEMENTS

I wish to thank Prof. A.E. van Wyk for his academic

assistance, the University of Pretoria and the National

Botanical Institute for putting their amenities at my

disposal, Mrs S. Perold for the SEM micrographs and Mrs

A. Romanowski for her photographic skills, Dr D.J.B.

Killick for the Latin diagnosis and Miss A. Pienaar for

the line drawings.

REFERENCES

PIENAAR, B.J. 1991. A new species of Vigna Savi (Fabaceae) from

southern Africa. South African Journal of Botany 57: 314—318.

PIENAAR, B.J. 1992. A taxonomic revision of the genus Vigna Savi

(Fabaceae) in southern Africa. M.Sc. thesis. Department of

Botany, University of Pretoria.

B.J. PIENAAR*

* Present address: P.O. Box 44057, Linden, Johannesburg 2104.

MS. received: 1992-08-17.

ASTERACEAE

AN EVALUATION OF HUTCHINSON’S BEETLE- DAISY’ HYPOTHESIS

INTRODUCTION

Some Cape Asteraceae species have conspicuous dark

markings on their ray florets. Such markings are usually

interpreted as ‘guides’ of various sorts (e.g. Faegri & Van

der Pijl 1979). However, Hutchinson (1946) suggested that

the dark raised marks on the ray florets of Gorteria diffusa

Thunb. mimicked herbivorous beetles burrowing head

down in the inflorescences. He noted that this species

appeared to have few beetle visitors and to suffer less

herbivory than other Asteraceae (such as an Arctotis sp.)

growing nearby. He hypothesised that the marks repelled

the beetles. In his review of plant mimicry worldwide,

Wiens (1978) considered this an exceptionally intriguing

example of Batesian mimicry. Despite this there still

appears to be a dearth of information on the interaction

between beetle-daisies and beetles. The purpose of this

note is to extend the concept of beetle -daisies and to test

Hutchinson’s hypothesis.

The beetles which commonly burrow into daisy flowers

are known as monkey-beetles (Coleoptera: Scarabaeidae:

Rutelinae/Hopliinae; Scholtz & Holm 1985). The sub-

family to which they belong is largely centred in the Cape.

Little is known about their ecology against which to test

Hutchinson’s hypothesis. The situation regarding the

relative absence of monkey-beetles on Gorteria diffusa

observed 45 years ago by Hutchinson remains unchanged

today (pers. obs. in Nieuwoudtville District). I did not see

any hopliinid visitors on this species in many hours of

observation in the spring of 1990. Since the hopliinids are

a large group and they visit many other plant species

(Whitehead, Giliomee & Rebelo 1987; pers. obs.),

Hutchinson’s hypothesis may be of more general relevance.

Assuming this, I studied the interaction of Aretotheca

calendula L. (Cape weed), a weedy daisy without dark

markings on the ray florets, and the beetle Heterochelus

sexlineatus Thunb., a herbivorous species with strong

cutting mandibles. This plant species is visited by many

hopliinid species (Scott & Way 1990) and I observed

Heterochelus sexlineatus visiting at least seven other plant

species at the study site, suggesting that there is only a

diffuse relationship between the two study taxa.

In Gorteria diffusa , the so-called beetle-daisy, the

‘beetle’ mark is a dark raised bump on the ray floret with

a white spot in the middle and with yellow ‘legs’. Under

ultraviolet light this ‘beetle’ does not appear significantly

different (pers. obs.). The number of ‘beetles’ per inflores-

cence is very variable (from none to a full ring with marks

on all ray florets) within and between individuals (pers.

obs.). The ‘beetles’ on inflorescences with a full ring

appear to be the least derived condition because they are

poorly differentiated and are similar in appearance to many

other Asteraceae with a ring of conspicuous dark basal

markings on ray florets (e.g. Gazania lichtensteinii). Even

Arctotis species (e.g. A. gumbletonii Hook, f.), which

Hutchinson (1946) suggested suffer more predation, have

complex dark basal markings on the ray florets.

It is thus possible that any dark marks near the base of

the ray floret or darkening of the disc found in other genera

such as Osteospermum, Dimorphotheca and Ursinia may

be mimicking beetles. In the Still Bay area, J. Vlok and

I noted an Ursinia species (close to U. paleacea (L.)

Moench) which also appears to be a beetle-daisy. In this

Bothalia 23,1 (1993)

71

TABLE 1. Number of Heterochelus sexlineatus beetles on manipulated and unmanipulated inflorescences of Arctotheca calendula. For plots

1 and 2 extra beetles were released (see text)

species some of the ray florets have been lost and through

the gaps they have left, large dark involucral bracts appear.

This exceptional modification is equally impressive in the

field as that of Gorteria diffusa. Ursinia is placed in the

tribe Anthemideae whereas all the other genera mentioned

are in the Arctotideae, indicating strong floral convergence.

According to K. Bremer (pers. com.) these types of dark

markings are probably restricted to the Cape Asteraceae.

If all the above modifications are shown to be a response

to monkey-beetles then I estimate that about 30 Cape

species could display the ‘beetle-daisy’ syndrome (see

Midgley 1991 for photographs of most of above examples).

The following information was collected to test Hutchin-

son’s hypothesis; (i) do numbers of beetle visitors differ

between inflorescences with artificial beetle marks and

those without, (ii) do numbers of beetles on unmanipu-

lated inflorescences suggest that the presence of one beetle

deters others?

MATERIAL AND METHODS

The behaviour of the hopliinid Heterochelus sexlinea-

tus was observed on a large ( > 500 individuals) popula-

tion of Arctotheca calendula growing wild in an arboretum

at Saasveld, near George in the southern Cape.

Sampling took place on warm days between 15h00 and

16h00 during October 1990. Inflorescences were manipu-

lated by marking the ray florets with brown, yellow and

black dots using commercial Artline pens. The yellow

marks were not visible (to human eyes) on the yellow ray

florets and thus served as controls to determine any other

non-visual effects of the marks on beetles. Numbers of dots

ranged from two to five and were approximately the same

size as the beetle. To simulate the Ursinia type model

(described above), from three to five ray florets were

removed from a sample of inflorescences. Inflorescences

were checked the following day for numbers of beetles.

In some cases inflorescences became unsuitable subse-

quent to marking and this accounts for unequal numbers

on Table 1. Because the numbers of beetles per inflores-

cence are low (less than 5% — see Results and conclusions)

in some instances beetles were captured from other areas

and released in the vicinity of study plots.

RESULTS AND CONCLUSIONS

Surveys indicated that 10 out of 200, six out of 100 and

nine out of 300 Arctotheca inflorescences had beetle visi-

tors (mean of less than 5% of inflorescences had visitors).

Ursinia anthemoides (L.) Poir. in the vicinity had less than

2% of inflorescences with visitors. The manipulation

experiments indicate that this beetle is virtually indifferent

to markings on the ray florets and to the absence of ray

florets (Table 1). The fact that considerable aggregation

of beetles occurs on inflorescences (e.g. up to eight

individuals in Table 2) suggests that the presence of an

individual is not inimical to others. Approximately twice

as many male beetles as females were found (Table 2).

Relative to the number of inflorescences, the beetles, and

especially the females, are rare. Consequently males

probably visit many inflorescences searching for mates.

The males fight for access to females (Midgley 1992). This

suggests that this beetle would be an effective pollinator

(its hairy body is often covered with pollen) but a rela-

tively insignificant herbivore (a few florets in a few

inflorescences in a population are damaged). The results

concerning manipulated inflorescences suggest that floral

markings have no negative effect on visitation. It is possible

that beetles are actually attracted to the marked florets.

However, on discovering that there are no real beetles on

the florets, they fly off. It was not, however, possible to

observe each visitor as it arrived on all manipulated in-

florescences simultaneously to see whether this was the

case. Although the hopliinid considered in this study is

a herbivore with strong cutting mandibles, many other

hopliininds are merely pollen feeders (Peringuey 1902).

It would make little sense for mimicry to evolve to repel

the non-herbivorous, pollen-carrying hopliinid beetles.

Also there would be little reason for an evolutionary trend

towards reduction in the number of beetle marks, if their

function is repulsion.

TABLE 2. — Distribution by gender of Heterochelus sexlineatus beetles

(M = male, F = female) in three samples of inflorescences of

which each had at least one beetle visitor

72

Bothalia 23,1 (1993)

The above (no repulsion, aggregation and feeding habits)

suggest that Hutchinson’s hypothesis is not complete. If

the markings act as mimics then it is probably to attract

beetles, presumably for their role in pollination. This

would then be a case of reproductive mimicry (sensu

Wiens 1978), similar in a way to pseudocopula tory orchids.

Obviously this study of one beetle and daisy species needs

to be broadened before Hutchinson’s intriguing hypothe-

sis of this little-studied syndrome of Cape plants is fully

tested.

ACKNOWLEDGEMENTS

I thank J. Vlok for field assistance, R. Oberprieler for

identifying the beetle and K. Bremer for his views on

Asteraceae markings.

REFERENCES

FAEGRI, K. & VAN DER PUL, R. 1979. Principles of pollination

biology. Pergamon Press, Oxford.

HUTCHINSON, J. 1946. A botanist in South Africa. Crawthom, London.

MIDGLEY, J.J. 1991. Beetle-daisies and daisy beetles. African Wildlife

45: 318, 319.

MIDGLEY, J.J. 1992. Why do some hopliinids have large hind legs?

Journal of the Entomological Society of Southern Africa 55:

157-163.

PERINGUEY, L. 1900-1902. Descriptive catalogue of the Coleoptera

of South Africa (Lucanidae and Scarabaeidae). Transactions of

the South African Philosophical Society 12: 1—220.

SCHOLTZ, C.H. & HOLM. 'E. 1985. Insects of southern Africa.

Butterworths, Durban.

SCOTT, J.K. & WAY, M .J. 1990. A survey in South Africa for potential

biological control agents against capeweed, Arctotheca calendu-

la (L.) Levyns (Asteraceae). Plant Protection Quarterly 5: 31-34.

WHITEHEAD, V.B., GILIOMEE, J.H. & REBELO, A.G. 1987. Insect

pollination in the Cape flora. In A. Rebelo, A preliminary

synthesis of pollination biology in the Cape flora. S.A.N.S.P. 141.

WIENS, D. 1978. Mimicry in plants. Evolutionary Biology 11: 365—403.

J.J. MIDGLEY*

* CSIR, Division of Forest Science and Technology, Private Bag X5011,

Stellenbosch 7599.

MS. received: 1991-08-16.

OXALIDACEAE

A NEW SPECIES OF OXALIS FROM THE WESTERN CAPE

Oxalis oculifera E.G.H. Oliver, sp. nov. in Sectione

Latifoliolatis , in genere singularis propter tubum corollae

rubrum papillatum partem alteram violaceo-roseum

annulo mediano alba, foliola subpeltata subparallela

glauca.

Planta parva, 5—15 mm alta. Bulbus ovoideus, 8x6

mm, ferragineus, vaginis papyraceis tectus, vaginis in parte

inferiore diagonaliter secedentibus. Folia 1—9; petiolus

2—20 mm longus, sparse glandulopilosus, roseus; foliola

3, interdum 2 vel 1, subparallela, 3.0— 4.5 x 3.0 mm, semi-

conduplicata, oblique subpeltata, basaliter subinfundi-

buliformia, late elliptica ad obovata, interdum oblongo-

elliptica, apice late obtusa vel plus minusve emarginata

base obtusa, glauca, adaxiale dense papillata, abaxiale

glabra sed interdum locis parvis croce is callosis, margine

hyalino et intra zona crocea gracili callosa; petioluli

0.3— 0.6 mm longi. Pedunculi uniflori, 20—37 mm longi,

erecti demum prostrati, sparse glandulopilosi, rosei;

bracteae absentes interdum 1 vel 2 in parte superiore,

filiformes vel lineares, ad 0.6 mm longae, sparse glandu-

lopilosae. Sepala 1.7— 1.9 x 0.7-0.9 mm, appressa, ovata

ad late ovata, interioria angustissima, subacuta, rasilia et

sparse glandulopilosa, zona marginali atropurpurea et zona

proxime interiore aurantiaca callosa, parte cetera viridi

erabescenti. Corolla ± 9—10 mm longa, late salviformis,

violaceorosea fauce cum annulo albo et tubo vinaceorabro;

petala 5—6 mm lata, oblique obovata ad late subspa thulata,

base param conjuncta, abaxiale sparsissime glandulopi-

losa, ecallosa, adaxiale in zona rubra papillata. Stamina

in seriebus tribus, base longitudine 0.5 -0.8 mm con-

juncta; antherae albae marginibus atropurpureis; fila-

menta purpurea sparse glandulopilosa; pollen tricol-

poratum, ellipsoideum, in antheris superioribus medi-

anisque album, in antheris inferioribus luteum. Ovarium

0.7— 1.0 mm longum, ovoideum, uniovulatum, in parte

superiore sparse glandulopilosum; styli in seriebus tribus.

mediani superioresque erecti ad param patentes, inferiores

valde porrecti, sparse glandulopilosi purpurei; stigmata

fimbriata, superiora medianaque purpurea, inferiora alba.

Figura 6.

TYPE. — 3118 (Vanrhynsdorp): Cape Province, Vanrhyns-

dorp Dist., Gifberg/Matsikamma area, central plateau

W of van Taakskom near top of the pass, 595 m,

(-DD), 12-06-1990, Oliver 9558 (STE, holotype; BOL,

K, PRE isotypes).

Dwarf stemless plants 5—15 mm high, aggregated into

clumps. Bulb ovoid, 8x6 mm, light reddish brown,

covered with papery sheaths splitting diagonally in lower

part. Rhizome vertical up to 20 mm long. Stem none or

very short up to 4mm long. Leaves 1—9 per plant; petiole

2—20 mm long, sparsely glandular pilose, pinkish; leaflets

mostly 3, occasionally 2 or 1, subparallel, 3.0— 4.5 x 3.0

mm, semiconduplicate, obliquely subpeltate, basally

subinfundibuliform, broadly elliptic to obovate, occasion-

ally oblong-elliptic, apically broadly rounded or slightly

emarginate basally rounded, glaucous, adaxially densely

papillate, abaxially glabrous, with hyaline margin and

inside this a thin orange callose zone, sometimes with

scattered small orange callose patches abaxially; petiolule

0.3— 0.6 mm long. Peduncle 1-flowered, 20—37 mm long,

erect becoming prostrate, markedly glandular pilose when

young and short, becoming sparsely so when mature,

pinkish; bracts usually absent, sometimes 1 or 2 on upper

part of peduncle, filiform or linear up to 0.6 mm long,

sparsely glandular pilose. Sepals 1.7— 1.9 x 0.7— 0.9 mm,

adpressed to corolla and joined at base, lobes ovate to

broadly ovate with inner ones narrowest, subacute,

glabrous and sparsely glandular pilose, with very dark

purple marginal zone and orange callose zone just inside

that, the rest green becoming reddish. Corolla about 9-10

mm long, broadly salver- shaped, violet-pink with white

FIGURE 6. — Oxalis oculifera : A, plant, X 2; B, leaf, adaxial view, on left, & abaxial view, on right, x 12; C, calyx, X 25; D, petal, abaxial

view, x 25; E, the three trimorphic series of the androecium and gynoecium, X 25; F, gynoecium, x 25. All drawn from the type, Oliver

9558 (STE).

I ring in throat and wine-red tube; petals 5-6 mm wide,

obliquely obovate to broadly subspathulate, joined slightly

at base, very sparsely glandular pilose abaxially on outer

margin, ecallose, papillate adaxially on red zone. Stamens

in 3 series, 2 series per plant, the shortest 1.2 mm long,

the middle 2.0— 2.3 mm long, the longest 3.0— 3.5 mm

long, all joined for 0.5— 0.8 mm at base, the uppermost

manifest, the rest included; anthers white with dark purple

edges; filaments purplish, sparsely glandular pilose; pollen

tricolporate, ellipsoid, in upper and middle anthers white,

in bottom anthers yellow. Ovary 0.7— 1.0 mm long, ovoid,

sparsely glandular pilose in upper half, 1-ovuled; styles

in 3 series, the lowest 0.5 mm long, markedly spreading

and white, the middle 1.2 mm long and the uppermost

series 3 mm long, the latter two series erect to slightly

spreading purplish sparsely glandular pilose; stigmas

fimbriate, the uppermost and middle series purple, the

lower series white. Figure 6.

This new species of Oxalis is remarkable in the genus

on account of the colour of its corolla and the distinctive

shape of its leaves. The corolla tube is wine-red with a

white ring in the throat, the rest of the corolla being violet-

pink. The glaucous leaflets are unusual for being obliquely

peltate with the petiolule attached abaxially to the lamina.

The leaflets are subparallel next to each other and not

spreading. They are semiconduplicate in a broad V-shape

with the basal portion above the petiole often partially

funnel-shaped.

O. oculifera falls within the Section Latifoliolatae which

Salter (1944) described as an unnatural assemblage of

species. It is defined as having those species forming

acaulescent plants with broad, but never linear or oblong,

leaves. Within the section this species is probably most

closely allied to O. petiolulata Bol. f.

O. petiolulata appears to be confined to the central and

northern parts of the Cedarberg near Clanwilliam where

it is known from only three collections, all in BOL ( H .

Bolus 8952, the type; Adamson in BOL 20492 and Po-

cock s.n.). All three have very differently sized leaves with

Adamson in BOL 20492 having the smallest leaves most

similar to those in the new species. The Pocock specimen

has no flowers and was determined with a query by Salter.

In O. petiolulata the plants are very sparsely and

minutely glandular pilose, not glabrous as recorded by

Salter. The sepals are sparsely ciliate with short rather

thick hairs. The leaflets have about six distinct black

callose ‘lumps’ towards the edges and nothing sub-

marginally, whereas in the new species the callose dots

are small, orange in colour and are randomly scattered

and there is a distinct, very thin, submarginal orange

callose line. In O. petiolulata the callose region on the

sepals is black and confined to two large patches near the

apex, whereas in the new species the callose region is

orange and submarginal down the sepal just below the thin

marginal black zone.

The bulbs in O. oculifera are quite different from those

in O. petiolulata. In the former the bulb is small and ovoid,

the tunics splitting with diagonal slits like an inverted V,

whereas in the latter they are narrowly ovoid and attenuate

with longitudinally splitting tunics.

The leaflets in the new species are very distinctive. They

are subparallel, adpressed and partially conduplicate. The

glaucous-grey colour of the leaves is very striking in the

living material. This is produced by numerous translucent,

shortly papillate cells on the upper surface. The remark-

able feature is the position of the point of attachment of

the leaflet to the petiolule. In O. petiolulata the petiolule

74

Bothalia 23,1 (1993)

is attached at the base of the lamina. In O. oculifera the

petiole is attached about one quarter of the way along the

undersurface of the lamina which is thus semipeltate. In

some leaflets the two sides of the basal portion of the

lamina are a little more adnate producing a slightly funnel-

shaped base to the leaflet.

Salter (1944) noted that he had not seen any living

material of O. petiolulata but described the flowers as ‘deep

pink with a purple eye in the throat, tube dull yellow’. F.

Bolus (1918) described the corolla from his father’s

collection as ‘purpurea’ and Adamson gave ‘deep pink with

a dark throat’ on his label. This would indicate that the

corolla was tricoloured with a purple ring at the mouth

of the tube which is yellow, quite unlike the very distinc-

tive wine-red tube forming the eye in O. oculifera. Salter

(1935) when describing his O. oculata (= O. callosa R.

Knuth) gave the colours as ‘roseo-rubra..tubo luteo, in

faucibus purpureo-annulata’ and noted that the dark purple

‘eye’ in the corolla mouth is a character which he found

in some forms of other species, but in O. callosa it

appeared to be always present. His drawing which was

repeated under O. callosa (Salter 1944), shows the dark

‘eye’ as a ring in the mouth of the corolla.

This feature of the red tube was mentioned to Salter

when I made the first collection in 1965. I told him I had

noted that all the flowers in the population possessed the

striking red eye. He remarked that this was unknown to

him in the genus and that the material constituted a distinct

new species.

O. oculifera is confined to one major population and

several nearby smaller populations in a small area on the

summit of the mountain plateau between the Gifberg and

Matsikamma Mountains (Figure 7). I have not been able

to locate any other populations, which could very well

be found on this extensive range with more intensive

searching. The plants occur in shallow gritty sand on

quartzitic sheet rock of the Table Mountain Group. The

main population grows where water seeps over the rock,

in some places as a small streamlet, the plants standing

in the shallow water. The plants are dominant, forming

pure stands of many individuals or clumps which are a

remarkable sight when in full bloom in May and early

June, a month after the first autumn rains.

This contrasts with the spring flowering period,

September/October, for O. petiolulata for the Bolus and

Pocock collections. Surprisingly the Adamson collection

was collected in flower in May 1933. Pocock recorded her

collection as ‘growing in marshy places and slowly flowing

streams’.

Salter (1944) noted that all the native South African

species are trimorphic. He mentioned that overseas

workers had recorded that the pollen in Oxalis was of

different sizes. Salter himself had noted that the colour

of the pollen varied but that he had not recorded the details

for the South African species. O. oculifera is trimorphic

within the populations studied and possesses two distinct

pollen types (Figure 6E). White larger grains are produced

by the middle and uppermost anthers and yellow smaller

grains by the lowest set of anthers. This feature raises the

intriguing problem of pollen selection in the pollination

FIGURE 7. — The known distribution of Oxalis oculifera.

syndrome of the species. In the wild no insect visitors other

than the occasional thrips crawling around in the flower,

were observed.

The material studied for this species included living

material of Oliver 9558 flowering in cultivation at Stellen-

bosch during May 1992.

Specimens examined

O. oculifera

CAPE.— 3118 (Vanrhynsdorp): Vanrhynsdorp Dist., Gifberg/Matsikam-

ma area, central plateau W of Van Taakskom near summit of the pass,

595 m, (-DD), 10-06-1965, Oliver in STE 32028 (PRE, STE, MO);

ibid., 12-06-1990, Oliver 9555 (BOL, K, PRE, STE).

O. petiolulata

CAPE. — 3219 (Wuppertal); Clanwilliam, Cedarberg, Krakadouw,

1 000 m, (— AA), 3-05-1933, Adamson in BOL 20492 (BOL); near

Wuppertal and Krakadouw, 1 000 m, (— AA/AC), 8-10-1897, H. Bolus

8952 (BOL, holo.); Cedarberg, between Middelberg hut and Crystal Pool,

1 220-1 525 m, (-AC), 09-1930, Pocock s.n. (BOL).

REFERENCES

BOLUS, F. 1918. Novitates africanae. Annals of the Bolus Herbarium

1: 19-28.

SALTER, T.M. 1935. Plantae novae africanae. Journal of South African

Botany 1: 129—144.

SALTER, T.M. 1944. The genus Oxalis in South Africa. Journal of South

African Botany , Suppl. Vol. 1: 1—355.

E.G.H. OLIVER*

* Stellenbosch Herbarium, P.O Box 471, Stellenbosch 7599.

MS. received: 1992-06-09.

Bothalia 23,1 (1993)

75

PTERIDOPHYTA— ADIANTACEAE

A NEW CYTOTYPE FOR ACROSTICHUM AUREUM

Acrostichum is a genus with a pantropical distribution

and consists of three, or perhaps more, species. A. aureum

L. is the most widespread of these and also has a

pantropical distribution.

On the west coast of Africa A. aureum occurs from

Guinea-Bissau to northern Angola and on the east coast

from Zanzibar to the Transkei. A. aureum is largely an

estuarine species which is commonly associated with the

upper edge of mangrove communities. It generally grows

in brackish or saline flats of alluvial mud or muddy sands.

Although it is predominantly an estuarine species it occurs

sporadically at considerable distances from the coast. In

Zimbabwe it was found at an elevation of approximately

500 m at a hot spring with salt efflorescences (Jacobsen

1983). Plants have also been recorded 100 km from the

coast in Saudi Arabia, 823 m above sea level, where it grows

in a slightly saline alkaline seepage (Collenette 1991).

Cytological observations in A. aureum have been carried

out on plants from Sri Lanka (Manton & Sledge 1954),

Ghana ( Manton 1959), southern India (Abraham et al.

1962) and Jamaica (Walker 1966). In all events the chromo-

some number reported was 2n = 60.

A collection of A. aureum made by me, Roux 1945

(NBG), in a seasonally marshy area on the banks of the

St Lucia Estuary, on the northern Natal coast, have now

been studied cytologically. Acetocarmine preparations of

root tips taken from this cultivated plant revealed a

chromosome number of 2n = 120 (Figure 8). This plant

may therefore be considered a tetraploid. The suggestion

made by Tryon & Tryon (1982) that polyploidy, which is

a common phenomenon in tropical pteridophytes, never

developed in the genus, can therefore no longer be up-

held. Most chromosomes are of the acrocentric and

telocentric type with few being submetacentric.

* *

v

„ s*.

^ %

A

&

JR* K

\

1

/H

tl

V

FIGURE 8. — Root tip mitosis of

Acrostichum aureum showing

120 chromosomes, Roux 1945

(NBG).

REFERENCES

ABRAHAM, A., NINAN, C.A. & MATHEW, P.M. 1962. Studies on

the cytology and phylogeny of the pteridophytes. VII. Observa-

tions on one hundred species of south Indian ferns. Journal of

the Indian Botanical Society 41: 339—421.

COLLENETTE, I.S. 1991. Acrostichum aureum: an inland record from

Saudi Arabia. Fern Gazette 14: 57.

JACOBSEN, W.B.G. 1983. The ferns and fern allies of southern Africa.

Butterworths, Durban, Pretoria.

MANTON, I. 1959. Cytological information on the ferns of west tropical

Africa. In A.H.G. Alston, The ferns and fern allies of west tropical

Africa: 75—81.

MANTON, I. & SLEDGE, W.A. 1954. Observations on the cytology

and taxonomy of the pteridophyte flora of Ceylon. Philosophi-

cal Transactions of the Royal Society of London, B, 238: 127—185.

TRYON, R.M. & TRYON, A.F. 1982. Ferns and allied plants with

special reference to tropical America. Springer-Verlag, New York,

Heidelberg, Berlin.

WALKER, T.G. 1966. A cytological survey of the pteridophytes of

Jamaica. Transactions of the Royal Society, Edinburgh 66:

169-237.

J.P ROUX*

* Compton Herbarium, National Botanical Institute, Private Bag X7,

Claremont 7735.

MS. received: 1992-05-12.

76

Bothalia 23,1 (1993)

FIGURE 9.—Argyrolobium rotundi folium, Edwards & Ackermann 329 (NU). A, habit; B, leaf; C, fruit; D, standard, frontal view; E, keel; F,

wing; G, calyx; H, androecium. Scale bars: A = 10 mm; B = 5 mm; C = 4 mm; D-F, H = 2 mm; G = 4 mm.

Bothalia 23,1 (1993)

77

FABACEAE

NOTES ON THE GENUS ARGYROLOBIUM (CROTA LAR1EAE) INCLUDING A NEW SPECIES FROM SOUTHERN AFRICA

Argyrolobium rupestre (E. Mey.) Walp. comprises

‘small slender-stemmed few-flowered prostrate or shortly

ascending plants from mountainous areas the length of

eastern Africa’ (Polhill 1968). Tropical representatives of

A. rupestre have been ‘loosely subdivided’ into four

subspecies by Polhill (l.c. ). The complex is notoriously

difficult to classify due to the poor resolution of characters

and Polhill (l.c.) concedes that ‘some of the forms may

possibly have evolved separately from more robust local

species but whatever their origin only slight differences

in habit and indumentum distinguish them’. Whereas his

revision covered high altitude specimens from South

Africa, no mention was made of the lowland element

discussed here and routinely included within A. rupestre.

Argyrolobium rotundifolium T. J. Edwards, sp.

nov., A. rupestri affinis sed habitu prostrato; foliolis sub-

orbiculatis cum pilis rufo-tomentosis; corolla alis sine

sculptura differt.

TYPE. — Natal, 2930 (Pietermaritzburg): between Peace-

vale & Drummond, (— DD), Edwards & Ackermann 329

(NU, holo. ; K, E, PRE, iso.).

Herbaceous perennial, up to 150 mm tall, but usually

prostrate, well branched, stems shortly rufous-tomentose,

becoming glabrous. Leaves moderately tomentose above,

densely tomentose beneath; leaflets broadly ovate to sub-

orbicular 14-32 x 10—25 mm, petiole tomentose, 10-30

mm long; apex obtuse to rounded, apiculate; margins

densely rufous-tomentose; stipules free, setaceous to

lanceolate, up to 10 x 1.5 mm. Inflorescence pseudo-

umbellate, 1— 4(— 6)-flowered, initially terminal but

becoming leaf-opposed; peduncle (30-)40— 150 mm long;

bracts linear or narrowly elliptic, up to 6 x 1 mm,

bracteoles linear, up to 3 mm long. Flowers dimorphic.

Calyx sparsely to densely pilose; upper lobes 8—10 mm

long, lower lip 8—10 mm long, lobes 3—4 mm long.

Corolla bright yellow becoming russet; standard sub-

orbicular, 11-14 x 12—14 mm, adaxial surface sericeous,

base cordate, claw 1.0— 1.5 mm long; wings obovate, 9—12

x 4.5 -6.0 mm, distally pilose, without sculpturing, claw

1.5— 2.0 mm long; keel cymbidiform, 8—10 x 4.5— 5.5

mm, pilose on lower margin, claw 1.5— 2.5 mm. Stamens

monadelphous, sheath fused above. Pistil narrowly oblong;

ovary 6-8 x 1-2 mm; style 3-4 mm long. Fruit rufous-

pilose, compressed, up to 45 x 5 mm. Seed subglobose,

laterally compressed, 2.5— 3.0 mm in diameter, yellow to

light brown, hilar rim raised.

Floral dimensions in the description refer to chasmoga-

mous flowers. Cleistogamous flowers are of no use in

species delimitation.

A. rotundifolium (Figure 9) is allied to A. rupestre but

is distinguished by its consistently prostrate habit (A.

rupestre is better described as ascending or decumbent),

broadly elliptical to orbicular leaflets, shortly rufous-

tomentose indumentum and the lack of sculpturing on

its wing petals. This species is predominantly coastal in

distribution whereas A. rupestre is limited to inland areas

(Figure 10). In southern Natal A. rupestre approaches the

coast on the high lying areas around Kokstad and Harding,

however, no contact zones with A. rotundifolium have been

observed.

NATAL. — 2731 (Louwsburg): Itala Nature Reserve, (— CA), Germis-

huizen 2232 (PRE). 2831 (Nkandla): Ngoye, (-DC), Huntley 852 (NU).

2832 (Mtubatuba): Hluhluwe Game Reserve, Mbhombe, (— AA),

Hitchins 598 (NH, NU). 2931 (Stanger): Nonoti Lagoon, (-AD), Ward

9121 (PRE). 3030 (Port Shepstone): Oribi Gorge, Riverview, (— CA),

Van Wyk 5064 (PRE). 3130 (Port Edward): 7 km from Port Edward on

the Izingolweni Rd, (-AA), Hilliard 1738 (NU).

CAPE. — 3228 (Butterworth): The Haven, (—BA), Gordon-Gray 557

(GRA). 3229 (Talemofu): Hole in the wall, (— AA), Germishuizen 1860

(PRE).

Argyrolobium lotoides Harv. in Flora capensis 2:

595 (1862). Type: Transkei, H. Bowker 366 ( TCD, holo.!;

K!, PRE! iso.). Figure 11.

A. variopile N.E. Br. : 18 (1906); B-E. van Wyk: 395 (1987) synon.

nov. Syntypes: Natal, Volksrust, hillside near Charlestown, Wood 5693

(BOL!, K!, NH!, PRE!), Wood 6355 (K!, NH!, SAM!).

A. hirsuticaule Harms: 179 (1917) synon. nov. Type: Transkei. Zuur-

bergen, Schlechter 6571 (Bf, holo.; BOL, lecto.! selected here).

A. leptocladum Harms: 180 (1917) synon. nov. Type: Kokstad, around

Clydesdale, Tyson 1256 pro parte (Bf, holo.; BM! , icono. selected here).

A. thodei Harms: 184 (1917) synon. nov. Type: Witzieshoek, Thode

20 (Bf, holo.; BM!, icono. selected here).

Lotononis magnistipulata Diimmer: 299 (1913). Type: Faku’s Territory

(probably Transkei), Sutherland s.n. (K, holo.).

Early collections of this species, made by Drege, bear

the name Chasmosyne pilosissima but no published

description was traced. The three names published by

Harms were based on differences in vestiture and leaf

dimensions. These highly variable characters form a

continuum and are thus unsuitable for delimiting species.

The specimens from which A. leptocladum and A. hir-

suticaule were described were destroyed in Berlin. The

drawings of these specimens at BM are therefore selected

FIGURE 10. —The known distribution of Argyrolobium rotundifolium,

#; and A. rupestre, A. in southern Africa.

78

Bothalia 23,1 (1993)

FIGURE 11. — A rgyrolobium lotoides, Browning 183 (NU). A, habit; B, dissected calyx, inner surface; C, standard, adaxial surface; D,

wing; E, keel; F, androecium. Scale bars: A = 10 mm; B— F = I mm.

Bothalia 23,1 (1993)

79

as iconotypes. The type of A. leptocladum is part of

a mixed collection {Tyson 1256)\ unfortunately only

specimens of A. amplexicaule remain from this gathering

(these are listed below under additional specimens

examined).

The species (Figure 12) occurs commonly in highland

sourveld and is frequently associated with rocky outcrops.

TRANSVAAL. — 2630 (Carolina): Chrissiemeer, (—AC), Theron 2383

(PRE). 2729 (Volksrust): hillside near Charlestown, (— BD), Wood 5693

(BOL, K, NH, PRE), Wood 6355 (K, NH, SAM).

ORANGE FREE STATE.— 2828 (Bethlehem): Generaalskop, (-DA),

Roberts 3298 (PRE).

NATAL. — 2730 (Vryheid): Naauwhoek, (—AD), Devenish 2124 (E,

K, NU, PRE, S). 2929 (Underberg): Sani Pass, (-CB), Hilliard & Bum

15534 (E, NU, PRE); Browning 183 (NU). 3029 (Kokstad): Zuurberg

near Weza, (— BC), Hilliard & Bum 8070 (E, MO, NU, PRE).

LESOTHO. — 2927 (Maseru): Molmo Ntuse mountain road, (— BD),

Schmitz 7248 (PRE). 2928 (Marakabei): Semonkong, (— CC), Jacot

Guillarmod 1812 (PRE). 3027 (Lady Grey): Ben McDhui, (— DB), Gal-

pin 6607 (PRE).

CAPE.— 3028 (Matatiele): Quachasnek, hillside facing Maluti Hotel,

(—BA), Gordon-Gray 4042 (E, NU).

Additional specimens examined

Argyrolobium amplexicaule

NATAL. — 3029 (Kokstad): in hills around Clydesdale, (— BD), Tyson

1256 (BM, BOL, K, SAM, UPS).

ACKNOWLEDGEMENTS

My gratitude is extended to the Natal University

Research Fund for financial assistance. The cited herbaria

are thanked for the loan of specimens and Mr M. Lambert

is thanked for translating the Latin diagnosis.

REFERENCES

BROWN, N.E. 1906. II. Diagnoses Africanae, XIV. Kew Bulletin 1906:

15-30.

DUMMER, R. A. 1913. A synopsis of species of Lotononis Eckl. &

Zeyh., and Pleiospora Harv. Transactions of the Royal Society

of South Africa 3 : 275 —335.

HARMS, H. 1917. Weitere Beobachtungen iiber Kleistogamie bei

affikanischen Arten der Gattung Argyrolobium. Berichte der Deut-

schen Botanischen Gesellschaft 35: 175—186.

HARVEY, W.H. 1862. Leguminosae. Flora capensis , Vol. 2. Hodges

Smith, Dublin.

POLHILL, R.M. 1968. Argyrolobium Eckl. & Zeyh. (Leguminosae) in

tropical Africa. Kew Bulletin 22: 145—168.

VAN WYK, B-E. 1987. Taxonomic notes on Argyrolobium variopile

(Fabaceae), and the status of Lotononis magnistipulata. South

African Journal of Botany 53 : 395 - 397.

T.J. EDWARDS*

* UN/FRD Unit for Plant Growth and Development, Botany Department,

University of Natal, P.O. Box 375, Pietermaritzburg 3200.

MS. received: 1992-03-12.

ALLISONIACEAE

THE HEPATIC, CALYC U LARI A CRISPULA (M ETZGERIALES ) REPORTED FROM MALAWI AND ZAMBIA

Calycularia crispula Mitt. , in Journal of the Proceed-

ings of the Linnean Society of Botany 5: 122 (1861); Pande

& Udar: 331 (1956); Jones: 497 (1985). Type: Himalayas,

Sikkim, J.D. Hooker 1679 (G, iso.).

Calycularia golae Gerola: 471-485 (1947). Type: Abyssinia, Vondo,

Valova s.n.

Thallus procumbent, dark bottle-green, to olive-green,

nearly translucent, delicate and easily torn when wet;

almost colourless in dead basal parts, with brownish

margins when dry; in overlying mats, on tree bark or on

soil covering tree boles; firmly adherent to substrate along

midrib throughout its length; 7.5—23.0 mm long, simple

or with a single terminal bifurcation (Figures 13A; 14A),

branches 5.0— 7.5 x 7.5— 9.0 mm, occasionally short lateral

branches arising ventrally near base, basal part semi-

lobate; wings thin and wide, margins slightly scalloped

and strongly undulate, otherwise entire; apex notched

(Figure 13E) with the 2 terminal lobes overlapping

somewhat; ventral scales situated between lobes and visi-

ble from above, arched over edge, small and inconspicu-

ous, mostly confined to apices of ventral face, colourless,

up to 500 pm long, 3 or 4 cell rows wide at base, central

cell row longer and tapering to uniseriate tip, lateral rows

shorter, all ending in and laterally with one to several slime

papillae, ± 50 pm, with spherical top (Figure DC); rhi-

zoids very long and dense, smooth, faintly brown, ± 20

pm in diameter, restricted to midrib ventrally; midrib

± J/5 of branch width, ± 280 pm (or more) thick and

generally consisting of 12— 15 rows of thin-walled homoge-

neous cells (Figure 13D), in cross section 4—6-sided, ±

42.5 x 27.5—32.5 jam, in longitudinal section ± rectangu-

lar, up to 137 pi n long, without a central strand of con-

ducting cells, gradually becoming thinner laterally and

wings eventually unistratose (Figure DD); epidermal cells

of midrib slightly thicker-walled, rectangular to pentago-

nal, generally 55 x 25 pm, in cross section 32.5 x 25.0—

32.5 ^m, containing numerous chloroplasts, ± 5.0 pm

80

Bothalia 23,1 (1993)

FIGURE 13. — Calycularia crispula. A, male thallus with androecia; B, scale-like involucre which individually covers antheridium; C, ventral

scale with slime papillae; D, part of transverse section of thallus, showing one unistratose wing and costa without central conducting strand;

E, ventral face of apex of thallus, showing apical notch and slime papillae; F, chloroplasts and small oil bodies inside thallus cells, seen

from above; G, cells along margin of thallus seen from above. A— C, F— G, S.M. Perold 2677 ; D, S.M. Perold 2668. A, X 9.5; B, X

90; C, x 125; D, E, x 45; F, x 400; G, x 150.

%,w

Bothalia 23,1 (1993)

81

FIGURE 14, — Calycularia crispula. A, female thallus with young sporophyte enclosed in perianth; B, close-up of sporophyte; C, longitudinal

section through capsule; D, involucral scales; E, cluster of archegonia protected by involucral scales, near apex of thallus; F, calyptra showing

unfertilized archegonium at base; G, perianth with laciniate mouth; H. transverse section of capsule wall, rod-like thickenings indicated

by arrows; I, cells in fragment of capsule wall, thickenings from above seen as dots. A— I, S.M. Perold 2668. A, x 6.5; B, C, X 12;

D, x 90; E, x 20; F, G, x 18; H, I, x 300.

82

Bothalia 23,1 (1993)

in diameter, as well as small oil bodies (Figure 13F);

cells in wings, from above, pentagonal or hexagonal,

45-50 x 30—50 /xm; cells at margin rectangular,

32.5—57.5 x 17.5—22.5 /am or pentagonal, 25 x 30 /xm

(Figure 13G).

Dioicous. Androecia consisting of 3 or 4(5) crowded,

± parallel rows of male scales overlying the midrib in the

body of the thallus, reduced to 2 rows in the terminal

branches (Figure 13A), antheridia short-stalked, globular,

± 250 x 240 /xm when mature, each one covered by a

forwardly directed laciniate scale (Figure 13B, C), laciniae

up to 550 /xm long x 250 (im wide at base, composed

of cells 45 —75 x 45 /x m. Gynoecia acropetally arranged

and singly placed dorsally, near the notched apex and when

present, above the furcation of the branches (Figure 14A,

B), slightly raised on a swelling, each with a group of

± 6—11 young archegonia (Figure 14E), ± 325 /xm long,

the necks composed of 6 cell rows and the tips brownish;

just posterior to the archegonia and at their sides subtended

by an involucre of narrow, laciniate scales (Figure 14D),

erect to somewhat forwardly directed; when mature, the

capsule spherical to ovate, leaning obliquely forward, 1750

x 1500 /x m, containing a mass of spores and elaters and

raised on the seta, here only + 1000 ix m long (Figure 14C)

and not maximally elongated; capsule wall brown,

bistratose (Figure 14H), cells square or rectangular, 37.5

X 37.5 to 75 X 20 /x m, cell walls thin, but with some

conspicuous orange-coloured semi-annular bands or

stiffening rods, the latter projecting into the cell lumen

(Figure 141); perianth obconic and large when mature,

enclosing capsule, green, with laciniate mouth (Figure

14G), 3100 ix m long including + 600 /xm long laciniae,

cells ± 75.0 x 37.5 /xm; calyptra delicate and transparent

(Figure 14F), bistratose, cells mostly pentagonal, ± 50.5

x 37.5 n m, with unfertilized archegonia attached to base.

Spores with distal face rounded (Figure 15C, D), proximal

face flat or indented (Figure 15A), light brown, trans-

lucent, diameter 35.0—42.5 /xm, densely echinate, spines

occasionally rather blunt, straight or bent (Figure 15B),

7.5 ix m long and 5.0 /xm wide at base, in between them

small projections arising at corners of shallow areolae.

Elaters bispiral (Figures 15F; 16), 10.0 /xm thick, ± 310

/xm long, tips solid, rounded. Vegetative reproduction by

stipe arising from group of cells in colourless, seemingly

dead marginal or basal parts of thallus, almost lunate, 175

x 375 ix m, soon narrowly winged; also by ventral branches

arising from midrib.

Grows on tree trunks and apparently also on soil, in

sheltered, damp places in montane forest.

DISCUSSION

Calycularia crispula (family Allison iaceae, order Metz-

geriales) is well known in the Himalayas. An excellent

description and line drawings of Indian material were

given by Pande & Udar (1956), who also studied the

ontogeny of the gametangia and the embryogenesis. Jones

(1985) remarked that his records of C. crispula were only

the second report of this species from Africa [Tanzania

and Malawi (which includes part of Chowo Forest)], the

first was by Gerola (1947), when he described C. golae

from Ethiopia (= Abyssinia). Jones believed that this

species should be treated as a synonym of C. crispula.

Arnell (1960) had misidentified a specimen of C. golae ,

collected by Dr John Eriksson in Ethiopia (= Abyssinia),

as Monocolea gottschei Lindb. (Grolle 1985), which does

not, however, grow in Africa. Besides the Himalayas

(and Indomalaya) the genus Calycularia is also known

from Taiwan, Japan, Siberia, Alaska (Schuster 1982), the

Aleutian Islands and Pacific northwest of North America

(Davidson & Smith 1992), as well as Mexico (Grolle

1980), and it is probably of Laurasian origin. Although

FIGURE 15. — Calycularia crispula. SEM micrographs of spores. A, ?proximal face of spore; B, spore seen from side; C, D, distal face ol spore;

E, part of spore much enlarged; F, young spores and elaters. A— E, S.M. Perold 2668', F, S.M. Perolil 2677. A-D, x 995; E, X 1990;

F, X 260. Micrographs by S.M. Perold.

Bothalia 23,1 (1993)

83

A-.'t '

5

FIGURE 16. — Calycularia crispula, LM photograph of elater, S.M.

Perold 2677, x 350.

Stephani (1900) had assigned six species to the genus, it

now appears to have just two, with C. crispula and C. laxa

Lindb. & Arnell [confined to the polar regions of Alaska

and Siberia (Davidson & Smith 1992)], the only species

belonging here, because Schuster (1982) transferred C.

radiculosa to the new monotypic genus, Sandeothallus

Schust.

The specimens collected by me are appreciably smaller

than the thallus sizes reported by Jones (1985), i.e. 20—50

mm long. My collections must be young plants, as only

one mature sporangium was found, which had not yet

dehisced. Pande & Udar (1956) reported it as dehiscing

by 4—6 irregular valves. An elaterophore was not observed

either; apparently it is not well developed in this species.

The seta did not elongate much and the sporangium

remained inside the perianth; with maturity the seta

reportedly elongates and raises the sporangium above the

perianth. The antheridia may not always be in continuous

rows and may grow in groups. The small ventral scales

are mostly confined to the region close to the apices of

the plants examined; their presence distinguishes

Calycularia from Pallavicinia lyelli and Dumortiera

hirsuta, which lack them. The SEM spore micrographs

taken in this study compare well with those of Inoue &

Hibino (1984).

Specimens examined

ZAMBIA. — 1033: Nyika Nat. Park, (—DA), Chowo Forest, on bark

of tree next to path, 15-4-1991, S.M. Perold 2668.

MALAWI.— 1033: Nyika Nat. Park, (-DA), 15 km W of Chilinda,

in forest on Parinari excelsa tree, 16-4-1991, S.M. Perold 2677.

ACKNOWLEDGEMENTS

I wish to express my sincere gratitude to Dr R. Grolle,

Jena, for kindly identifying my two collections and

Dr Grolle and Prof. T. Poes, Eger, for refereeing the

manuscript. My thanks also to the typist, Mrs J. Mulvenna,

and to the photographer, Mrs A. Romanowski, (both of

NBI) for their valuable contributions.

REFERENCES

ARNELL, S. 1960. Hepatics collected by Dr John Eriksson in Abyssinia

in 1958. Svensk Botanisk Tidskrift 54: 187—192.

DAVIDSON, P.G. & SMITH, D.K. 1992. Calycularia crispula (Hepati-

cae) in the Aleutian Islands and Pacific northwest of North

America. The Bryologist 95: 266—269.

GEROLA, F.M. 1947. Epatiche dell’ Abissinia meridionale (Raccolte

dalla Missione Ittiologica in A. O.I. Lavori di Botanica. Istituto

Botanico dell’ Universita Padova 3: 471—485.

GROLLE, R. 1980. Miscellanea hepaticologica 201—210. Journal of

Bryology 11: 325—334.

GROLLE, R. 1985. Miscellanea hepaticologica 231—240. Journal of the

Hattori Botanical Laboratory 58: 197—202.

INOUE, H. & HIBINO, R. 1984. Studies on spore morphology of

hepatics (1). Metzgeriales. Bulletin of the National Science

Museum (Tokyo). Ser. B (Botany) 10: 177—189.

JONES, E.W. 1985. African hepatics XXXV. Some new or little-known

species and some noteworthy records. Journal of Bryology 13:

497-508.

MITTEN, W. 1861. Hepaticae of the East Indies. Journal of the Proceed-

ings of the Linnean Society 5: 89—128.

PANDE, S.K. & UDAR, R. 1956. Studies in Indian Metzgerineae. III.

Calycularia crispula Mitten. Phytomorphology 6: 331—346.

SCHUSTER, R. 1982. Studies on Hepaticae, LIX. On Sandeothallus

Schust., gen. n. and the classification of the Metzgeriales. Nova

Hedwigia 36: 1—16.

STEPHANI, F. 1900. Species hepaticarum. Vol. 1. Bulletin de I 'Herbier

Boissier 6: 300—411.

S.M. PEROLD

MS. received: 1992-03-16.

Bothalia 23,1: 85-90 (1993)

First report on the presence of Enterobryus species (Trichomycetes:

Eccrinales) in South Africa and the description of three new species

G.J.M.A. GORTER*

Keywords: Diplopoda, Eccrinales, Enterobryus spp. nov., gut fungi, millipedes. South Africa, Trichomycetes

ABSTRACT

The occurrence of Trichomycetes in the Republic of South Africa is reported for the first time. Thus far only various

juliform millipedes have been examined for the presence of these fungi. Three new species of Enterobryus have been found

and are described in detail, viz. E. centroboli Gorter, E. chaleponci Gorter and E. zinophorae Gorter.

UITTREKSEL

Die voorkoms van Trichomycetes in die Republiek van Suid-Afrika word vir die eerste keer aangemeld. Tot dusver is

slegs verskillende wurmagtige duisendpote vir die aanwesigheid van hierdie swamme ondersoek. Drie nuwe spesies van

Enterobryus is gevind en word beskryf, nl. E. centroboli Gorter, E. chaleponci Gorter en E. zinophorae Gorter.

INTRODUCTION

Enterobryus species are gut fungi which belong to the

class Trichomycetes, lower fungi which are placed in

the Subdivision Zygomycotina (Lichtwardt 1986). All

Trichomycetes are characterised by the production of

sporangiospores and the presence of a secreted holdfast

with which the thalli are attached, mostly to the internal

cuticular surface of certain Arthropoda. Four orders

have been recognised in the Trichomycetes by Manier

& Lichtwardt (1968), viz. Harpellales, Asellariales,

Eccrinales and Amoebidiales. Sexual reproduction has

been observed only in the Harpellales and in one genus

(Enteropogon) of the Eccrinales (Hawksworth etal. 1983).

None of these fungi have been grown in axenic culture

except a few of the Harpellales ( Smittium spp. and

Trichozygospora chiron omidarum) and one of the

Amoebidiales ( Amoebium parasiticum) (Lichtwardt 1978).

However, this does not imply that they are parasites. Their

relationship with their hosts is considered to be commen-

salistic, the host neither benefitting nor suffering from the

association (Moss 1979). They produce one or more types

of sporangiospores basipetally. Lichtwardt (1954, 1958)

designated these spore types as A, B, C, D, E, F and G.

They are attached by a holdfast to the hindgut cuticle of

Diplopoda, less often Insecta (Coleoptera) and Crustaceae

(Lichtwardt 1986). As noted by Lichtwardt (1978), the

Eccrinales is, from a taxonomic viewpoint, the most

difficult order of the Trichomycetes to study, probably on

account of the unstable morphology of its members and

the inability of researchers to culture them.

Work on the Eccrinales has been done principally in

Europe, particularly in France (Duboscq et al. 1948;

Manier 1950, 1954, 1961, 1963, 1964, 1969, 1970; Pois-

son 1927, 1928, 1929, 1931) and in the United States of

America (Leidy 1849a, b; Lichtwardt 1954, 1957a, b, 1958,

1960a, b), but studies have also been done in India

(Rajagopalan 1967) and Japan (Lichtwardt et al. 1987).

* Plant Protection Research Institute, Private Bag X134, Pretoria 0001.

MS. received: 1992-02-11.

This article records the presence of these fungi for the

first time in South Africa.

MATERIALS AND METHODS

Fungi for study were obtained from live Diplopoda. The

procedures for dissection of the millipedes and observa-

tion for the presence of fungi were similar to those used

by Lichtwardt (1954). However, to facilitate the cutting off

of the head and anal segment, the live millipedes were first

immobilised by placing them in tubes of a suitable dimen-

sion and cooling them in a freezer for 10 or more minutes,

depending on the size of the millipede. The gut was then

removed by grasping the posterior portion of the hindgut

with a forceps and gently pulling the gut from the body.

Next, the gut was cut open with finely pointed, curved

scissors after which the undigested gut content was re-

moved with the aid of a thin jet of water. As the removal

of the cuticle with the fungi from the gut was found to

be laborious and not always successful, most examinations

were made of the fungi in situ, either in water or after

treatment with lactophenol, with or without cotton blue.

Observations on nuclei were made after fixing in Clarke’s

fluid (Bradbury 1973) and staining with Heidenhain’s iron-

alum-haematoxylin.

Millipedes were obtained from various parts of the

country. Studies were confined to the juliform type of

millipedes.

RESULTS AND DESCRIPTIONS

During the rainy seasons of 1989/90 and 1990/91, dozens

of millipedes of different species including common ones

such as Doratogonus setosus (Voges), D. circulus (Attems)

and Orthoporoides pyrocephalus (L. Koch), were exa-

mined for the presence of Trichomycetes. Every species

showed some signs of infestation. However, most of them

lacked primary spores whereas the mycelium with secon-

dary spores had traits similar to those described species

which produce primary spores. This has made identifica-

tion impossible. Nevertheless, in different groups of

86

Bothalia 23,1 (1993)

millipedes, three fungi were encountered which differed

sufficiently from described species to be recognised as

new.

Figures illustrating the newly described species serve

as holotypes because the slides on which they are based

are subject to deterioration.

Enterobryus centroboli Gorter, sp. nov.

Hyphae rectae vel undulatae, 3000-3800 x 7.5—12.5

(—15) ^m, plerumque hyalinae. Cytoplasma granulosum

et obscurum in hyphis laterioribus. Hyphae saepe tumidae

prope basim sed angustiores ad retinaculum. Retinaculi

orbis basalis 10.0— 12. 5(— 15) pm diam., caulis perbrevis,

circa 5.0 x 7.5 pm, vel absens. Sporangiosporae (typus A)

15.0 x 7.5 jtm (uninucleatae) et 40 60 x 7.5 ^tm (quadrt-

nucleatae). Sporae multinucleatae (typus E) 90—125 x

10-20 pm. Sporae uninucleatae (typus F), 5.0-7.5 x 10

pm.

Habitat affixa in intestino posteriore Centroboli spp. ,

Natal, 1991.

TYPUS. — Figura 1A— F nititur laminis vitreis PREM

50874 (NC 20) , PREM 50875 (NC 18) et PREM 50876 (NC

43) Centroboli lawrencei (Schubart).

The fungus develops unbranched thalli which are

attached to the anterior part of the hindgut, just below but

occasionally also just inside the sphincter muscle (the

muscle which divides the fore- and hindgut). Hyphae also

develop towards the posterior end of the hindgut. In both

cases, the hyphae are fairly straight to undulating, 7.5—12.5

(—15) ^m wide, those at the front are up to 3800 pm long

and those at the back up to 3000 pm. Most hyphae are

hyaline but some of those in front tend to become wider

and more opaque as the result of dense granular cytoplasm

towards the hyphal apex. The narrower hyaline hyphae

produce at their apex short or longer type A sporangio-

spores. The short ones (15.0 x 7.5 pm) are uninucleate

and the longer ones (40—60 x 7.5 /rm) are quadrinucleate

(Figure 1A, C). The wider, more opaque hyphae produce

3 to 4 apical or sometimes intercalary spores, 90—125 x

10—20 pm (Figure IF), which appear to be multinucleate

and of Lichtwardt’s type E.

Holdfasts are produced at the straight basal ends of the

hyphae which are often swollen (by 2.5— 5.0 ^im) towards

the holdfast and then narrow towards the point of attach-

ment (Figure IE). The holdfast has a very short stalk which

is often shorter than wide, e.g. 5.0 x 7.5 ^m. The diameter

of the attachment disc is comparatively small, viz.

10.0-12. 5(— 15) pm.

FIGURE 1. — Enterobryus centroboli Gorter. A, left, hyphae with 4-nucleate A spores, right, germinated type A spore fastened to substrate; B,

empty sporangia (arrowed) from which type A spores have escaped; C, germinated type A spores which immediately develop holdfasts

(arrowed); D, stacks of uninucleate primary spores; E, base of hypha which widens towards holdfast; F, type E spores. Scale bars = 25 fim.

Bothalia 23,1 (1993)

87

FIGURE 2. — Enterobryus zinophorae Gorter. A, rounded base of folded spore inside basal part of sporangium; B, apex of developing folded

spore; C, spiral base of hy pha devoid of sporangia; D, mature folded spore inside sporangium in straight part of hypha (spore tip arrowed);

E, holdfast with short stalk (arrowed) on base of straight hypha; F, empty sporangium cell with transverse folds in wall. Scale bars = 25 pm.

Hyphae which produce primary sporangiospores are

rare or absent. These hyphae are found at the posterior

end of the hindgut where they produce stacks of up to 85

uninucleate spores towards their apex (type F) which are

slightly shorter than wide, viz. 5.0— 7.5 x 10 pm (Figure

ID). Here, too, the holdfast has a short stalk, up to 5 pm,

with a small disc diameter of 10.0—12.5 pm.

Habitat: attached to the hindgut lining of Centrobolus

spp. from Natal (Cedara and Mtunzini), 1991.

TYPE. — Figure 1A— F, based on slides PREM 50874

(author’s collection number NC 20), PREM 50875 (NC

18) & PREM 50876 (NC 43) all of Centrobolus lawrencei

(Schubart).

Enterobryus zinophorae Gorter, sp. nov.

Hyphae rectae, flexae vel undulatae, 2000—3000 x

(7.5— )10.0— 12. 5(— 15) pm, raro incurvatae prope basim.

Cytoplasma hyalinum. Hyphae leviter dilatatae ad basim.

Retinaculum cum caule brevissimo, breviore quam lato,

haud longiore quam 6 pm: orbis basalis 12.5—20.0 ^m

diam. Nonnullae hyphae 1—6 cellulas sporangiales

175—300 x 10 pm ad apicem producentes. Sporangia

matura sporam unam longissimam replicatamque

continentia. Sporae basi tumida et rotunda sine palliolo

apicali. Post liberationem sporarum parietes sporangiales

contrahuntur fiuntque multirugosi. Sporae liberatae,

250—350 x 7.5—10 /xm, a (undo germinantes, hyphas

breves cum retinaculo producentes. Formas alias spora-

rum non vidi.

Habitat affixa intra sphincterem intestini Zinophorae

spp.

TYPUS. — Figura 2 A— F nititur laminis vitreis PREM

50877 (MP 4) et PREM 50878 (MP 7) Zinophorae

diplodontae (Attems), Pretoria, Transvaal, Dec. 1990.

Bothalia 23,1 (1993)

The thalli are inside the sphincter muscle. The hyphae

are hyaline, straight, bent or undulating towards their apex

but occasionally hooped near their base (Figure 2C),

2000-3000 x (7.5— )10.0— 12.5(— 15) /mi. Hyphae have

rounded ends and may widen slightly towards their base

where they secrete a holdfast with a short stalk not longer

than 6 /im and usually shorter than wide (Figure 2E).

Some hyphae produce a number of cells 175-300 /cm long

and usually 10 /cm wide towards their apex. These are

sporangia which, when mature, contain long, folded spores

(Figure 2B, D) with a slightly swollen obtuse base (Figure

2A) and a pointed apex without a protecting cap as

illustrated by Manier et al. (1974: fig. IF). These spores

are 250-350 x 7.5-10.0 /cm. The sporangia, from which

they escape, contract to become empty cells with many

folds (Figure 2F). The escaped spores straighten out, im-

mediately develop a hypha from the rounded base and

E

attach themselves to the sphincter lining. The hypha keeps

on growing after which the spore presumably degenerates

(empty filaments at the end of comparatively short hyphae

have been noted). No other type of spore has been

observed.

Habitat: attached to the sphincter muscle lining of

Zinophora spp.

TYPE.— Figure 2A-F, based on slides PREM 50877

( author ’s collection number MP 4) & PREM 50878 (MP

7) of Zinophora diplodonta (Attems), Pretoria, Transvaal,

Dec. 1990.

Enterobryus chaleponci Gorter, sp. nov.

Hyphae hyalinae, satis rectae sed plerumque circulares

prope basim, circa 2500 x (10.0 — )12. 5 — 15.0(— 17.5) /cm,

in circulis plerumque latiores quam alibi; latitudo

FIGURE 3. — Enterobryus chaleponci Gorter. A, short sporangia with folded spores (tips arrowed) inside hoops of swollen hyphae; B, mature

spore inside sporangium in straight part of hypha (spore tip arrowed); C, holdfasts without stalks on straight basal end of hyphae; D, germinated,

straightened-out acicular spore (base arrowed); E, multi nucleate spore of type E; F, germinated multinucleate spore attached to substrate

(holdfast arrowed). Scale bars = 25 ^m.

Bothalia 23,1 (1993)

89

aliquando etiam variat in partibus rectis. Retinaculum sine

caule, vel cum caule perbrevi; orbis basalis (12.5— )L5— 20

pm diam. Sporae longissimae, 125—230 pm, sine palliolo

apicali, replicatae, in sporangiis intra circulos 70 — 85 x

15 pm, in partibus rectis 105 — 150 x 12.5 /xm. Fundus

basalis sporae truncatus cum levi depressione in centro.

Cytoplasma hypharum latiorum dense granulosum prope

apicem. Hyphae latae aliquando cellulas longas (ad 125

/xm) producentes endosporium singulum multinuclearem

continentes.

Habitat affixa intra sphincterem intestini Chaleponci

polechanci Kraus, Baltimore, Transvaal, Mai. 1991.

TYPUS. — Figura 3A-F nititur laminis vitreis PREM

50879 (TB 5) et PREM 50880 (TB 30).

The thalli are inside the sphincter muscle. The hyphae

are hyaline, fairly straight but strongly hooped lower down,

+ 2500 x (10— )12.5— 15.0(— 17.5) pm. The hoops are often

wider than the straight parts of the hyphae, 15—20 pm.

The latter are of variable width, widening by 2 pm and

then narrowing again. A holdfast (Figure 3C) without a

stalk or with a very short one, e.g. 2.5 pm, develops at

the base of a hypha. The disc diameter varies from

(12.5 — )15 — 20 pm. The thicker hyphae and those that

widen from bottom to top, have a dense granular cytoplasm

and often become slightly brown. Folded spores have been

observed in the hooped as well as the straight part of

hyphae (Figure 3A, B). In the first, sporangia are 70—85

x 15 /xm; in the latter, 105—150 x 12.5 pm. The spores

are 125—230 x 10.0—12.5 pm and have a truncated basal

end with a slight depression in the centre. After escaping

from the sporangia, the folded spores straighten, germinate

basally and soon attach themselves to the inside of the

sphincter muscle (Figure 3D). At the apex of a hypha with

dense cytoplasm, a long cylindrical sporangium (up to 125

/xm) is often formed (Figure 3E). These sporangia are

slightly swollen in comparison with the rest of the hyphae

and each contains a multinucleate endospore.

Habitat: attached to the sphincter muscle lining in the

gut of Chaleponcus polechancus (Kraus), Baltimore,

Transvaal, May 1991.

TYPE. — Figure 3A— F, based on slides PREM 50879

( author’s collection number TB 5) & PREM 50880

(TB 30).

DISCUSSION

Apart from reporting the occurrence of Trichomycetes

for the first time in South Africa, a most interesting result

of this investigation was the discovery of sporangia

containing long, folded, pointed spores in some of these

fungi. These have hitherto been found only in millipedes

from Central Africa by Manier et al. (1974/75). They

described a number of species in which acicular, folded

spores are formed but noted that they develop only in the

narrow hyphae. This led Lichtwardt (1986) to pose the

question of whether the thalli with long, folded spores

perhaps belong to one fungal species, whereas the wider

thalli with different spores belong to different fungi. The

latter supposition is apparently supported by our finding

that, in E. chaleponci, the two types of spores present

mostly occurred alone in different specimens of a batch

of 36 millipedes of this species from one locality. However,

the fact that these spore types were found together, albeit

infrequently, and the fact that narrow hyphae could enlarge

to become wider, point to a close relationship.

The suggestion that hyphae with long, folded spores all

belong to one species is probably not valid because I found

clear differences in size between the folded spores of

E. chaleponci and E. zinophorae. The shape of their basal

ends also differed slightly while they develop at a different

locus. In E. zinophorae they develop only in the straight

part of hyphae, while in E. chaleponci also within the

hooped part of hyphae. Furthermore, Manier et al.

(1974/75) observed that in some Eccrinales species the

apex of the folded spore is protected by a solid point devoid

of protoplasm, whereas in other species this protection

is absent. These differences all indicate that different

species are involved.

ACKNOWLEDGEMENTS

I wish to thank the following persons sincerely for the

collection of the millipedes examined: Mesdames Pat

Caldwell, Louise Schutte, Jeany Heinlein and Dr R. L.

Kluge from the Plant Protection Research Institute, and

Prof. Albert Eicker of the University of Pretoria, Mrs Irene

Robinson and Mr H. A. Robinson.

I am also much indebted to Dr Richard L. Hoffman of

the Virginia Museum of Natural History, USA, for iden-

tification of the millipedes. Thanks are also due to Mr Paul

Hasse of the University of Pretoria for checking the Latin

descriptions and to Miss Alice Baxter of our Institute for

reading the manuscript.

REFERENCES

BRADBURY, S. 1973. Peacock’s elementary microtechnique, 4th edn.

Edward Arnold, London.

DUBOSCQ, O., LEGER, L. & TUZET, O. 1948. Contribution a la

connaissance des Eccrinides: les Trichomycetes. Archives de

Zoologie Experimental et Generate 86: 29—144.

HAWKSWORTH, D.L., SUTTON, B.C. & AINSWORTH, G.C. 1983.

Ainsworth & Bisby's dictionary of the fungi, 7th edn: 387. Com-

monwealth Mycological Institute, Kew.

LEIDY, J. 1849a. Enterobrus, a new genus of Confervaceae. Proceedings

of the Academy of Natural Science of Philadelphia 4: 225—233.

LEIDY, J. 1849b. Remarks upon several new species of Endophyta,

Enterobrus spiralis and Enterobrus attenuatus, and a new species

of Gregarina. Proceedings of the Academy of Natural Science

of Philadelphia 4: 245.

LICHTWARDT, R.W. 1954. Three species of Eccrinales inhabiting the

hindguts of millipedes with comments on the Eccrinids as a group.

Mycologia 46: 564-585.

LICHTWARDT, R.W. 1957a. Enterobryus attenuatus from the Passalid

beetle. Mycologia 49: 463 —474.

LICHTWARDT, R.W. 1957b. An Enterobryus occurring in the millipede

Scytonotus granulatus (Say.). Mycologia 49: 734—739.

LICHTWARDT, R.W. 1958. An Enterobryus from the millipede Boraria

Carolina (Chamberlin). Mycologia 50: 550-561.

LICHTWARDT, R.W. 1960a. An Enterobryus (Eccrinales) in a common

greenhouse millipede. Mycologia 52: 248-254.

LICHTWARDT, R.W. 1960b. New species of Enterobryus from south-

eastern United States. Mycologia 52: 743—752.

LICHTWARDT, R.W. 1978. Taxonomic problems in Trichomycetes related

to their growth in arthropod guts. In C.V. Subramanian, Taxono-

my of the fungi, Part I: 109—114. University of Madras, India.

LICHTWARDT, R.W. 1986. The Trichomycetes . fungal associates of

arthropods. Springer Verlag, New York.

LICHTWARDT, R.W., KOBAYASI, Y. & INDOH, H. 1987. Trichomy-

cetes of Japan. Transactions of the Mycological Society of Japan

28: 359-412.

90

Bothalia 23,1 (1993)

MANIER, J.-F. 1950 (1951). Recherches sur les Trichomycetes. Annales

des Sciences Naturelles, Botanique Ser. 11, 11: 53—162.

MANIER, J.-F. 1954. Essais de culture des Eccrina flexilis Leger et

Duboscq, Trichomycetes endocommensaux des Glomeris margi-

nata Villers. Annales Parasitologie Humaine et Comparee (Paris)

29: 265-270.

MANIER, J.-F. 1961. Eccrinides des Crustac^s recoltes sur les cotes du

Finistfere. Cahiers Biologie Marine 2: 313—326.

MANIER, J.-F. 1963. Trichomycetes parasites d’Isopodes Oniscoidea.

Annales des Sciences Naturelles, Botanique 4: 557-577.

MANIER, J.-F. 1964. Nouvelle contribution a 1’etude de Trichomycetes

(Eccrinales parasites d’amphipodes). Annales des Sciences

Naturelles, Botanique S6r. 12, 5: 767—772.

MANIER, J.-F. 1969 (1970). Trichomycetes de France. Annales des

Sciences Naturelles, Botanique S6r. 12, 10: 565—672.

MANIER, J.-F. 1970. Sur la frequence de Trichomycetes Eccrinales dans

le proctodeum des Myriapodes Diplopodes. Bulletin du Museum

National d'Histoire Naturelle 41: 91-95.

MANIER, J.-F., GASC, C. & BOUIX, G. 1974 (1975). Sur quelques En-

terobryus (Trichomyctes Eccrinales) parasites de Myriapodes

Diplopodes du Sud-Dahomey. Bulletin de I'Institut Fondamental

d’Afrique Noire 36: 614 —641.

MANIER, J.-F. & LICHTWARDT, R.W. 1968 (1969). Revision de la

systematique des Trichomycetes. Annales des Sciences Naturelles,

Botanique S6r. 12, 9: 519-532?

MOSS, S.T. 1979. Commensalism of the Trichomycetes. In L.R. Batra,

Insect-fungus symbiosis : 175 -227. Allanfield, Osman, Montclair,

N.J.

POISSON, R. 1927. Sur une Eccrinide nouvelle: Taeniellopsis orchestriae,

nov. gen., nov. sp., Protophyte parasite du rectum de 1’ Orchestra

bottae M. Edw. (Crust. Amphipode), Son cycle evolutif. Compte

rendu hebdomadaire Seances de l Academie des Sciences Paris

185: 1328, 1329.

POISSON, R. 1928. Eccrinopsis mercieri n. sp. Eccrinide parasite du

rectum de VOniscus asellus L. Son cycle evolutif. Compte rendu

hebdomadaire Seances de l Academie des Sciences Paris 186:

1765-1767.

POISSON, R. 1929. Recherches sur quelques Eccrinides parasites de

Crustaces Amphipodes et Isopodes. Archives de Zoologie

Experimentale et Generate 69: 179—216.

POISSON, R. 1931. Recherches sur les Eccrinides. Deuxifeme contribu-

tion. Archives de Zoologie Experimentale et Generate 74: 53 —68.

RAJAGOPALAN, C. 1967. An Enterobryus (Trichomycetes, Eccrinales)

in a millipede. Current Science 36: 20—22.

Bothalia 23,1: 91-104 (1993)

Mycorrhizal status of plants growing in the Cape Floristic Region, South

Africa

N. ALLSOPP* and W. D. STOCK*

Keywords: cluster roots, Fynbos, mediterranean-type shrublands, mycorrhizas, root systems

ABSTRACT

A survey of the mycorrhizal status of plants growing in the Cape Floristic Region of South Africa was undertaken to assess

the range of mycorrhizal types and their dominance in species characteristic of this region. Records were obtained by ex-

amining the root systems of plants growing in three Cape lowland vegetation types, viz. West Coast Strandveld, West Coast

Renosterveld and Sand Plain Lowland Fynbos for mycorrhizas, as well as by collating literature records of mycorrhizas

on plants growing in the region. The mycorrhizal status of 332 species is listed, of which 251 species are new records.

Members of all the important families in this region have been examined. Mycorrhizal status appears to be associated mainly

with taxonomic position of the species. Extrapolating from these results, we conclude that 62% of the flora of the Cape

Floristic Region form vesicular-arbuscular mycorrhizas, 23% have no mycorrhizas, 8% are ericoid mycorrhizal, 2% form

orchid mycorrhizas, whereas the mycorrhizal status of 4% of the flora is unknown. There were no indigenous ectomycor-

rhizal species. The proportion of non-mycorrhizal species is high compared to other ecosystems. In particular, the lack of

mycorrhizas in several important perennial families in the Cape Floristic Region is unusual. The diversity of nutrient acquir-

ing adaptations, including the range of mycorrhizas and cluster roots in some non-mycorrhizal families, may promote co-

existence of plants in this species-rich region.

UITTREKSEL

’n Opname van die status van mikorisas by plante wat in die Kaapse Floristiese Streek van Suid-Affika voorkom, is ondemeem

om die omvang te bepaal van verskillende mikorisatipes en hulle oorheersing in spesies kenmerkend van hierdie streek.

Rekords is verkry deur die wortelsisteem van plante in drie laagliggende Kaapse veldtipes, naamlik Weskus-strandveld, Weskus-

renosterveld en Sandvlakte-fynbos, vir mikorisas te ondersoek. Rekords uit die literatuur van toepassing op mikorisas wat

op plante in hierdie streek groei, is ook in ag geneem. Die status van mikorisas by 332 spesies word vermeld, 251 daarvan

nuwe rekords. Lede van al die belangrike families in die streek is ondersoek. Die status van mikorisas is skynbaar

hoofsaaklik met die taksonomiese posisie van die spesie geassosieer. Ons het tot die gevolgtrekking gekom dat 62% van

die flora van die Kaapse Floristiese Streek vesikuler-arbuskulere mikorisas vorm, 23% geen mikorisas het nie, 8% erikoiede

mikorisas vorm, 2% orgidee-mikorisas het, terwyl die mikorisa-status van 4% van die flora onbekend is. Daar was geen

inheemse spesies wat ektomikorisas gevorm het nie. Die proporsie nie-mikorisaspesies is hoog in vergelyking met die wat

in ander ekosisteme aangetref word. Die afwesigheid van mikorisas by verskeie belangrike meeijarige families in die Kaapse

Floristiese Streek is veral ongewoon. Die verskeidenheid aanpassings om voedingstowwe te bekom, insluitend die omvang

van mikorisas en troswortels by sekere nie-mikorisafamilies, mag die gelyktydige bestaan van plante in hierdie spesieryke

streek bevorder.

INTRODUCTION

It is generally accepted that most terrestrial plants

probably form mycorrhizal associations between their roots

and certain fungi, although the vast majority of plants

growing in natural ecosystems have not had their mycor-

rhizal status confirmed (Trappe 1987; Newman & Reddell

1987). The mycorrhizal status of plants reflect both their

taxonomic affinities and their ecology. Investigations on

the mycorrhizal status of plants in various parts of the

world indicate that the major terrestrial biomes can be

characterized by specific mycorrhizal types (Read 1991).

Surveys of mycorrhizas show that trees of forests and

woodlands are either ectomycorrhizal or vesicular-

arbuscular mycorrhizal; herbaceaous plants and shrubs in

grasslands and shrublands usually form vesicular-

arbuscular mycorrhizas; boreal and temperate heathlands

are dominated by ericoid mycorrhizal species; and

disturbed ecosystems by non-mycorrhizal weed species

(see Brundrett 1991 for references). Whereas the mycor-

rhizal status of some floras is well documented (e.g. British

* Department of Botany, University of Cape Town, Private Bag,

Rondebosch 7700, South Africa.

MS. received: 1992-05-12.

Isles by Harley & Harley 1987), little is known about both

the mycorrhizal associations of plants in the Cape Floristic

Region (CFR) and their functional role in low nutrient

ecosystems.

The vegetation of the CFR contrasts sharply, in terms

of taxonomic composition and vegetation structure, with

the surrounding southern African vegetation. The Cape

flora has a high species diversity (± 8 500 species), around

68% species endemism (Bond & Goldblatt 1984), and high

beta and gamma species turnover (Cowling 1990). Agricul-

ture, urbanization and alien plant invasion are a severe

threat to this flora as a result of the limited range of many

plant species, and have led to the destruction of much of

the lowland vegetation (Hall 1983). Mycorrhizas act as soil

nutrient-absorbing organs for the plants. As such they will

influence the physiology of individuals, as well as their

interactions with other plants growing in the same

community (Harley 1989; Read 1991). Recognizing the

patterns of distribution and understanding the ecological

role of mycorrhizal types in a community may be crucial

to understanding the dynamics which shape plant com-

munities (Allsopp & Stock in press).

This study collates published records of the mycorrhizal

status of plants occurring in the CFR as defined by

92

Bothalia 23,1 (1993)

Bond & Goldblatt (1984). In addition, the mycorrhizal

status of plants growing in three lowland vegetation types,

West Coast Strandveld, West Coast Renosterveld and Sand

Plain Lowland Fynbos, is reported for the first time. The

aim of this paper is to provide information on the mycor-

rhizal status of plant species which may be of significance

in explaining vegetation patterns and plant functioning in

the CFR.

STUDY AREAS

Three study sites representing West Coast Strandveld,

at Melkbosstrand (33°45'S 18°27'E), West Coast Renoster-

veld on the hill, Joostenberg, on the farm Hercules Pillar

(33°46'S 18°46'E), and Sand Plain Lowland Fynbos at the

Fynbos Biome intensive study site (33°31S 18°32'E) at

Pella, were chosen to investigate the mycorrhizal status

of a broad range of plants growing in threatened habitats

in the CFR. The classification of the vegetation categories

follows that of Moll et al. (1984). The strandveld vegeta-

tion, growing on coarse/medium sandy soil (organic matter

2.2%, pH of 7.5, total P 422 ^g g-1 (Witkowski &

Mitchell 1987)) is a broad-leaved, sclerophyllous 1.0— 2.5

m high shrubland with a large succulent component

(Boucher 1983). The Renosterveld vegetation, growing on

a fine sand/clayey soil [organic matter 4.9% , pH 4.1, to-

tal P 127 ^ig g-1 (N. Allsopp unpublished)] is an ever-

green, cupressoid or microphyllous shrubland, 1-2 m

high, dominated by Elytropappus rhinocerotis with strong

Lowland Fynbos affinities (Tansley 1982; Boucher 1983).

The Sand Plain Lowland Fynbos growing on medium

textured sandy soil [organic matter 1.4— 3.4%, pH

4.6-4.8, total P 23-34 g-1 (Mitchell, Brown &

Jongens-Roberts 1984)] is an ericoid- leaved, sclerophyl-

lous vegetation, 0.75—1.5 m high with some taller shrubs,

characterized by the presence of Phylica cephalantha

(Boucher 1983). Vegetation surveys at the three sites have

recorded 56, 63 and 215 perennial species at the Strand-

veld (Siegfried 1981), Renosterveld (Tansley 1982) and

Lowland Fynbos (Boucher & Shepherd 1988) sites respec-

tively. In addition annuals and bulbous species are

numerically important components of all three vegetation

types (Boucher 1983; Bond & Goldblatt 1984).

MATERIALS AND METHODS

Root collection

Roots were collected during August, September and

early October when unthickened roots were common while

the soil was moist. Two collections were made at both the

Strandveld (during August 1987 and September 1989) and

Renosterveld sites (during October in 1988 and September

1989). The Lowland Fynbos site was sampled six times

over four years (June and August 1986, August and Sep-

tember 1987, September 1988, August 1989). Two 25 x

25 m plots were set up at a site on each collection day.

The plots at the Strandveld site were situated 60—300 m

inland of the high water mark. The Renosterveld plots were

on the NW-SW-facing lower slopes of Joostenberg. At the

Lowland Fynbos site, plots were randomly scattered

throughout the 269 ha study site.

Roots of one representative of all species occurring in

the plots were sampled. In addition, plant species not in

the plots, but encountered in the vicinity, were sampled.

Smaller plants, including annuals, perennial seedlings and

bulbous plants, were excavated with entire root systems.

Roots of larger shrubs were collected by carefully tracing

the root system from the main stem until young, unthick-

ened roots were encountered. However, for some species,

including members of the Anacardiaceae and Ebenaceae,

few young roots could be found despite extensive excava-

tion along roots down to 1 m. At the Renosterveld site

some species were not sampled because they grew only

in narrow cracks among rocks (e.g. Olea sp.). At the

Strandveld site the large size of dominant shrubs and

density of the vegetation at ground level, as well as

spininess of some species, precluded collection of these

species’ roots.

Young roots were removed from surrounding soil in the

field and immediately placed in vials containing 10% KOH

for clearing. Wherever possible, 500 mm of root per plant

was collected. Roots were cleared for one week at 20 °C,

and then rinsed under running tap water (Smith & Bowen

1979). Where necessary, pigmented roots were decolour-

ized with H2O2 or NaClO. This was followed by acidi-

fication in 1 M HC1 and staining in 0.05% Trypan blue

in lactic acid solution (Kormanik & McGraw 1982). Root

segments were permanently mounted in a polyvinyl acid

solution and inspected at 100 and 400 times magnifica-

tion with a light microscope for mycorrhizal structures.

Plants were classified according to Cronquist (1988) and

species names follow Gibbs Russell et al. (1985, 1987).

Literature survey

All known records of the mycorrhizal status of plants

in the CFR were consulted. Only those records which

reported the mycorrhizas of plants actually growing in the

CFR are listed here. Confirmation of infection status of

some species was undertaken by examining roots of plants

growing in soil from their natural habitats in pot culture.

RESULTS

Vesicular-arbuscular mycorrhizas (VAM), characterized

by the presence of arbuscules in the inner cortical cells

with or without vesicles (Harley & Smith 1983), were the

most common type of mycorrhiza (61% of species

examined) (Appendix). Infections regarded as vesicular-

arbuscular (VA) mycorrhizal but morphologically distinct

from the above types were formed by Aristea dichotoma,

which formed intracellular coils similar to those described

by Brundrett & Kendrick (1990) in Trillium grandiflorum,

while vesicular-arbuscular mycorrhizal fungi in Orphium

frutescens and Sebaea exacoides formed structures typical

of those seen in other members of the Gentianaceae

(Jacquelinet-Jeanmougin & Gianinazzi-Pearson 1983).

Infection formed by the ‘fine endophyte’ (Greenall 1963)

was occasionally seen, but was never exclusively found

on one species. Ericoid mycorrhizas (ERIC) were found

in the hair roots of all members of the Ericaceae examined

(Appendix). They are characterized by the formation of

coiled and branched, fine hyphae in the cortical cells (Read

1983). Orchid mycorrhizas (ORCH) were seen in the two

Disa spp. examined (Appendix) and consist of characteris-

tic coarse, coiled intracellular hyphae (Harley & Smith

1983). No ectomycorrhizal infection was seen in the in-

digenous species examined. Introduced ectomycorrhizal

Bothalia 23,1 (1993)

93

species such as pines, oaks, poplar and eucalypts form

ectomycorrhizas in the CFR but the ectomycorrhizal fun-

gi were in all likelihood introduced with imported saplings

(Van der Westhuizen & Eicker 1987).

Ninety-one of the 332 species reported formed no

mycorrhizas (Appendix). These were concentrated in the

Caryophyllidae and the families Brassicaceae, Cras-

sulaceae, Proteaceae, Santalaceae, Zygophyllaceae,

Restionaceae and Cyperaceae. Plant roots which contained

occasional vesicles but no arbuscules were regarded as

functionally non-mycorrhizal (Hirrel et al. 1978).

Some earlier studies (Laughton 1964; Low 1980) have

reported endophytic mycorrhizas (ENDO) as being present

but descriptions or illustrations do not indicate structures

which are typical of mycorrhizas as they are presently

delimited (Harley & Smith 1983). Non-mycorrhizal fungi

were fairly frequent in both mycorrhizal and non-

mycorrhizal roots examined for this study. Therefore,

reports of fungal infection as ‘endophytic mycorrhizas’

should be viewed with caution. The most common non-

mycorrhizal root inhabiting fungus was Olpidium sp.,

which forms cysts and zoosporangia (Sampson 1939),

which may be mistaken for VA mycorrhizal vesicles if care

is not taken. Unidentified hyphal fungi, including dark,

septate hyphal fungi forming microsclerotia (DSH) similar

to those described by Haselwandter & Read (1980) in

alpine vegetation, were also present. The non-mycorrhizal

roots of members of the Proteaceae have been shown to

support a fungal flora that is distinctly different to that

found in the non-rhizosphere soil (Allsopp et al. 1987).

Infection by Olpidium sp. and other fungi was particularly

heavy in the root systems of members of the Poaceae and

Scrophulariaceae where they could obscure infection by

VA mycorrhizal fungi (Appendix).

DISCUSSION

The mycorrhizal status of many of the taxa recorded here

has not previously been reported, as can be expected, given

the high levels of endemism and species radiation in the

CFR and the paucity of mycorrhizal studies in this region.

All the important families, as well as the twenty largest

genera in the CFR (Bond & Goldblatt 1984), now have

had some of their members examined for mycorrhizas.

The endemic Penaeaceae and near endemic Bruniaceae

have VA mycorrhizal species. Families which need further

investigation are the Anacardiaceae, Ebenaceae, Jun-

caceae, and Celastraceae, as well as the endemic families

Stilbaceae, Grubbiaceae, Roridulaceae, Retziaceae and

Geissolomaceae. The lowland vegetation types have been

well covered and generalizations regarding their mycor-

rhizas can now be made. However, the mycorrhizal status

of the vegetation of habitats such as forests, seasonally

waterlogged soils, limestone and mountain ecosystems are

less well catalogued.

The absence of ectomycorrhizas is a notable feature of

this flora. Ectomycorrhizal structures are reported in many

plants growing in arid regions of Australia which belong

to families and genera also present in the CFR (Warcup

1980; Warcup & McGee 1983; McGee 1986; Bellgard

1991). Ectomycorrhizas are also known to occur in the low

nutrient soils of the Australian mediterranean heathlands

(Chilvers & Pryor 1965; Brundrett & Abbott 1991). In

addition, ectomycorrhizas have been found on trees

growing in other African ecosystems (Redhead 1968;

Hogberg & Piearce 1986). However, shrub vegetation

growing on Kalahari sands adjacent to ectomycorrhizal

woodlands was exclusively vesicular- arbuscular mycor-

rhizal (Hogberg & Piearce 1986). The reasons for the

exclusion of ectomycorrhizas from the CFR are not clear,

although this can possibly be ascribed to the absence of

an organic surface horizon which is usually associated with

the presence of ectomycorrhizas (Read 1991), and to

frequent disturbance by fire. For instance, in Italian

mediterranean ecosystems on calcareous soils, canopy

cover values for ectomycorrhizal plant species nine years

after fire was a quarter of that in unbumt forest (Puppi

& Tartaglini 1991). However, these explanations do not ac-

count satisfactorily for their absence in the CFR, because

ectomycorrhizas occur in fire-prone communities in

Australia with low soil organic matter (Brundrett & Abbott

1991).

The explosive speciation that the genus Erica has

undergone in the CFR (± 530 spp.) implies that ericoid

mycorrhizas are unusually common in this area. Cowling

et al. (1990) have suggested that edaphic specialization of

the endophyte has powered this speciation, but, as yet,

supporting evidence is lacking. An interesting feature of

ericoid mycorrhizal plants in the mediterranean-climate

regions of the world is their co-existence with other plant

species, whereas in more temperate regions they usually

form almost pure stands in areas where soil degradation

has produced soil conditions which plant roots and other

mycorrhizas cannot tolerate (Leake et al. 1989).

All the non-mycorrhizal families in this study have been

reported as such before, although some have had very few

species examined for mycorrhizal colonization (Trappe

1987). Many of the non-mycorrhizal species in this study

fall into the Caryophyllidae which is roughly equivalent

to the Centrospermae (Cronquist 1988) which was origi-

nally regarded as non-mycorrhizal (Gerdemann 1968).

Subsequent studies have shown that many species in this

group are capable of forming mycorrhizas (Tester et al.

1987), and that some families are typically mycorrhizal,

e.g. Cactaceae (Miller 1979). However, despite these

exceptions, 80% of the species in the Caryophyllidae

which have been examined are either non-mycorrhizal or

facultatively mycorrhizal (Trappe 1987). Mechanisms

which enable some species to actively exclude mycorrhizal

fungi, when exposed to viable inoculum, are unclear

(Tester et al. 1987; Koide & Schreiner 1992).

In dicotyledonous species, weedy, herbaceous plants

often lack mycorrhizas or are weakly mycorrhizal (Malloch

et al. 1980; Trappe 1987) and it has been noted that some

species are less likely to form mycorrhizas when coloniz-

ing disturbed sites than adjacent undisturbed areas (Miller

1979; Reeves et al. 1979). In our study, the annuals in the

Scrophulariaceae were usually non-mycorrhizal, although

a few individuals form typical VAM.

Anaerobic conditions in waterlogged soils have been

invoked to explain the absence of mycorrhizas in some

plants (Anderson et al. 1984), and Tester et al. (1987)

advance this as an explanation of the absence of mycor-

rhizas in most of the Cyperaceae. In this study the mem-

bers of the Cyperaceae and the Restionaceae examined

94

Bothalia 23,1 (1993)

were non-mycorrhizal while growing in well -drained soil

with other mycorrhizal plants, although both families are

often associated with waterlogged conditions, and so this

does not seem to be the only reason for the exclusion of

mycorrhizas from these taxa. Although Powell (1975)

reported mycorrhizal structures in some roots of members

of the Cyperaceae, he concludes that they are functionally

non-mycorrhizal due to the possession of a fine root

system. This complements Baylis’ (1975) proposal that the

magnolioid root form with poorly developed root hairs

would be more strongly mycorrhizal than finer root

systems. Two important perennial families in the CFR,

which do not form mycorrhizas (viz. Proteaceae and

Restionaceae), are characterized by the formation of cluster

roots, the rootlets of which are densely covered in long

root hairs (Purnell 1960; Lamont 1972a, 1982). In addi-

tion, cluster roots have been observed on members

of the Cyperaceae (Lamont 1974), the genus Aspalathus

(Fabaceae) (this paper and M. Cocks unpublished data)

and Australian members of the Fabaceae (Lamont 1972b;

Brundrett & Abbott 1991) , which typically have low VAM

infection levels. The absence or low infection levels of

mycorrhizas in the taxa forming cluster roots support

Baylis’ (1975) proposition that mycorrhizas will be less

important when root systems are finer. The loss of the

ability to form mycorrhizas is regarded as an evolutionarily

advanced feature (Trappe 1987).

The mycorrhizal status of the species in the CFR seems

to be a reflection of their taxonomic position, although

Newman & Reddell (1987) warn that very few families

form exclusively one type of mycorrhiza or are consistently

without mycorrhizas. This can be expected when world-

wide the higher taxa of angiosperms are poorly correlated

with their ecological niches (Cronquist 1988). Life form

or environmental factors do not satisfactorily explain the

absence of mycorrhizas in longer lived plants such as

members of the Proteaceae, Restionaceae and Zygo-

phyllaceae in the CFR and this must be regarded as a

taxon-related characteristic for many groups. Reports of

VA mycorrhizal species among the Proteaceae in New

South Wales, Australia (Bellgard 1991) and ectomycorrhizal

Faurea saligna (Proteaceae) in Zambia (Hogberg &

Piearce 1986), indicate that the mycorrhizal status of mem-

bers of this family should be investigated with respect to

soil fertility, as mycorrhizas are absent in members of this

family growing in low nutrient soils of the CFR and

Western Australia (Brundrett & Abbott 1991). Members

of families such as the Aizoaceae and Mesembryan-

themaceae, which are commonly found associated with

disturbed areas in the CFR, are non-mycorrhizal when

growing in undisturbed ecosystems. This supports the

report that at the ecosystem level, patterns of mycorrhizal

and non-mycorrhizal species among weedy species

followed taxonomic divisions irrespective of growth form

(Pendleton & Smith 1983). As most of the data here are

obtained from plants growing in the field and mycorrhizal

status was usually consistent at the family level, generali-

zations can be made regarding the mycorrhizal status of

the Cape Flora, provided cognisance is taken that excep-

tions may arise. The mycorrhizal status of the flora of the

three study sites, representing three lowland vegetation

types is summarized in Table 1. If the mycorrhizal status

of species listed in Bond & Goldblatt (1984) is inferred

from that of taxonomically related species which have been

examined, we conclude that 62% of the flora form VAM,

TABLE 1. —Summary of the mycorrhizal status of the vegetation growing

in Sand Plain Lowland Fynbos, Renosterveld and Strandveld

communities, and the Cape Floristic Region (CFR)

VAM = vesicular-arbuscular mycorrhizal; ABS = non-mycorrhizal;

ERIC = ericoid mycorrhizal; ORCH = orchid mycorrhizal.

plants without mycorrhizas are the next largest group,

ericoid and orchid mycorrhizas are found in less than 10%

of the flora, and the mycorrhizal status of 4% of the flora

is unknown (Table 1).

The proportion of non-mycorrhizal species in the CFR

is high when compared to many other vegetation types

worldwide (Brundrett 1991). As non-mycorrhizal plants are

normally associated with high levels of disturbance, or

edaphically and climatically extreme conditions, the non-

mycorrhizal flora in the CFR is atypical in that represen-

tatives of two families that dominate the vegetation of the

CFR, the Proteaceae and Restionaceae, are non-

mycorrhizal. The evolutionary and ecological significance

of this needs further exploration. The diversity of mycor-

rhizal types is possibly an indication that no one type of

mycorrhiza or other nutrient acquiring adaptation is

pre-eminently suited to the environmental conditions in

the CFR and that the diversity of nutrient acquisition

mechanisms in the CFR has probably promoted species

co-existence.

ACKNOWLEDGEMENTS

We thank C. Allsopp, M. T. Deignan, E. C. February,

J. Jacobs, A. Jensen and H. Vaughan for assistance with

field work; J. Midgley and the nursery at Kirstenbosch

for providing plants; and A. Bean, E. Esterhuysen, C.

Johnson, H. P. Linder, P. Littlewort, J. Manning, F. Powrie

and E. van Jaarsveld for help with identifying plant species.

Mr S. de Wit is thanked for permission to collect roots

on his farm, Hercules Pillar.

REFERENCES

ALLSOPP, N., OLIVIER, D.L. & MITCHELL, D.T. 1987. Fungal popu-

lations associated with root systems of proteaceous seedlings at

a lowland fynbos site in South Africa. South African Journal of

Botany 53: 365—369.

ALLSOPP, N. & STOCK, W.D. in press. Density dependent interac-

tions between VA mycorrhizal fungi and even-aged seedlings of

two Fabaceae species. Oecologia.

ANDERSON, R.C., LIBERTA, A.E. & DICKMAN, L A. 1984. In-

teraction of vascular plants and vesicular-arbuscular mycorrhizal

fungi across a soil moisture-gradient. Oecologia 64: 111-117.

BAYLIS, G.T.S. 1975. The magnolioid mycorrhiza and mycotrophy in

root systems derived from it. In F.E. Sanders, B. Mosse & P.B.

Tinker, Endomycorrhizas : 373 —389. Academic Press, London.

BELLGARD, S.E. 1991. Mycorrhizal associations of plant species in

Hawksbury sandstone vegetation. Australian Journal of Botany

39: 357-364.

BOND, P. & GOLDBLATT, P. 1984. Plants of the Cape Flora. Journal

of South African Botany Suppl. Vol. 13.

Bothalia 23,1 (1993)

95

BOUCHER, C. 1983. Floristic and structural features of the coastal fore-

land vegetation south of the Berg River, western Cape Province,

South Africa. Bothalia 14: 669—674.

BOUCHER, C. & SHEPHERD, PA. 1988. Plant communities of the

Pella site. In M.L. Jarman, A description of the fynbos biome

intensive study site at Pella: 38—76. Occasional Report No. 33.

CSIR, Pretoria.

BRUNDRETT, M. 1991. Mycorrhizas in natural ecosystems. Advances

in Ecological Research 21: 171— 3D.

BRUNDRETT, M. & ABBOTT, L.K. 1991. Roots of Jarrah forest plants.

I. Mycorrhizal associations of shrubs and herbaceous plants. Aus-

tralian Journal of Botany 39: 445—457.

BRUNDRETT, M. & KENDRICK, B. 1990. The roots and mycor-

rhizas of herbaceous woodland plants n. Structural aspects of

morphology. New Phytologist 114: 469 —479.

CHILVERS, G.A. & PRYOR, L.D. 1965. The structure of eucalypt

mycorrhizas. Australian Journal of Botany D: 245-259.

COWLING, R.M. 1990. Diversity components in a species-rich area

of the Cape Floristic Region. Journal of Vegetation Science 1:

699-710.

COWLING, R.M., STRAKER, C.J. & DEIGNAN, M.T. 1990. Does

microsymbiont-host specificity determine plant species turnover

and speciation in Gondwanan shrublands? A hypothesis. South

African Journal of Science 86: 118-120.

CRONQUIST, A. 1988. The evolution and classification of flowering

plants. 2nd edn. The New York Botanical Garden, New York.

GERDEMANN, J.W. 1968. Vesicular-arbuscular mycorrhiza and plant

growth. Annual Review of Phytopathology 6 : 397-418.

GIBBS RUSSELL, G.E., REID, C., VAN ROOY, J. & SMOOK, L.

1985. List of species of southern African plants, edn 2, part 1.

Memoirs of the Botanical Survey of South Africa No. 51.

GIBBS RUSSELL, G.E., WELM AN , W.G. , RETIE F, E., IMMEL-

MAN, K. L., GERMISHUIZEN, G., PIENAAR, B.J., VAN

WYK, M., NICHOLAS, A., DE WET, C., MOGFORD, J.C.

& MULVENNA, J. 1987. List of species of southern African

plants, edn 2, part 2. Memoirs of the Botanical Survey of South

Africa No. 56.

GREENALL, J.M. 1963. The mycorrhizal endophytes of Griselinia

littoralis (Comaceae). New Zealand Journal of Botany 1:

389 -400.

HALL, A.V. 1983. Threatened plants of the southwestern comer of Africa.

Bothalia 14: 981-984.

HARLEY, J.L. 1989. The significance of mycorrhizas. Mycological

Research 92: 129— D9.

HARLEY, J.L. & HARLEY E.L. 1987. A checklist of mycorrhiza in

the British flora. New Phytologist 105: 1—102.

HARLEY, J.L. & SMITH, S.E 1983. Mycorrhizal symbiosis. Academic

Press, London.

HASELWANDTER, K. & READ, D.J. 1980. Fungal associations of roots

of dominant and sub-dominant plants in high-alpine vegetation

systems with special reference to mycorrhiza. Oecologia 45:

57-62.

HIRREL, M.C., MEHRAVARAN, H. & GERDEMANN, J.W. 1978.

Vesicular-arbuscular mycorrhizae in the Chenopodiaceae and

Cruciferae: do they occur? Canadian Journal of Botany 56:

28D-2817.

HOGBERG, P. & PIEARCE, G.D. 1986. Mycorrhizas in Zambian trees

in relation to host taxonomy, vegetation type and successional

patterns. Journal of Ecology 74 : 775 —785.

JACQUELINET-JEANMOUGIN, S. & GIANINAZZI-PEARSON, V.

1983. Endomycorrhizas in the Gentianaceae . I. The fungi

associated with Gentiana lutea L. New Phytologist 95: 663 —666.

KOIDE, R.T. & SCHREINER, R.P. 1992. Regulation of the vesicular-

arbuscular mycorrhizal symbiosis. Annual Review of Plant

Physiology and Plant Molecular Biology 43: 557—581.

KORMANIK, P.P. & MCGRAW, A.C. 1982. Quantification of vesicular-

arbuscular mycorrhizae in plant roots. In N .C. Schenck, Methods

and principles of mycorrhizal research: 37-46. The American

Phytopathological Society, Minnesota.

LAMONT, B. 1972a. The morphology and anatomy of proteoid roots

in the genus Hakea. Australian Journal of Botany 20: 155—174.

LAMONT, B. 1972b. Proteoid roots in the legume Viminaria juncea.

Search 3: 90.

LAMONT, B. 1974. The biology of dauciform roots in the sedge

Cyathochaete avenacea. New Phytologist 73 : 985 —996.

LAMONT, B. 1982. Mechanisms for enhancing nutrient uptake in plants

with particular reference to mediterranean South Africa and

Western Australia. Botanical Review 48: 597-689.

LAUGHTON, E.M. 1964. Occurrence of fungal hyphae in young roots

of South African indigenous plants. Botanical Gazette 125: 38—40.

LEAKE, J.R., SHAW, C. & READ, D.J. 1989. The role of ericoid mycor-

rhizas in the ecology of ericaceous plants. Agriculture, Ecosystems

and Environment 29: 237—250.

LOW, A.B. 1980. Preliminary observations on the specialized root mor-

phologies in plants of the western Cape Province. South African

Journal of Science 76: 5D— 516.

MALLOCH, D.W., PIROZYNSKI, K.A. & RAVEN, PH. 1980. Eco-

logical and evolutionary significance of mycorrhizal symbiosis

in vascular plants (a review). Proceedings of the National Academy

of Science, USA 77: 21D-2118.

MCGEE, P. 1986. Mycorrhizal associations of plants in a semi-arid com-

munity. Australian Journal of Botany 34: 385 -393.

MILLER, R.M. 1979. Some occurrences of vesicular-arbuscular mycor-

rhiza in natural and disturbed ecosystems of the Red Desert. Cana-

dian Journal of Botany 57: 619—623.

MITCHELL, D.T., BROWN, G. & JONGENS-ROBERTS, S.M. 1984.

Variation of the forms of phosphorus in the sandy soils of coastal

fynbos, southwestern Cape. Journal of Ecology 72 : 575 — 584.

MOLL, E.J., CAMPBELL, B.M., COWLING, R.M., BOSSI, L., JAR-

MAN, M.L. & BOUCHER, C. 1984. A description of major

vegetation categories in and adjacent to the fynbos biome. South

African National Scientific Programmes Report No. 83. CSIR,

Pretoria.

NEWMAN, E.L & REDDELL, P. 1987. The distribution of mycor-

rhizas among families of vascular plants. New Phytologist 106:

745 -751.

PENDLETON, R.L. & SMITH, B.N. 1983. Vesicular-arbuscular mycor-

rhizae of weedy and colonizer species at disturbed sites in Utah.

Oecologia 59: 296-301.

POWELL, C.L. 1975. Rushes and sedges are non-mycotrophic. Plant

and Soil 4: 481-484.

PUPPI, G. & TARTAGLINI, N. 1991. Mycorrhizal types in three mediter-

ranean communities affected by fire to different extents. Acta

Oecologica 12: 295—304.

PURNELL, H.M. 1960. Studies of the family Proteaceae. I. Anatomy

and morphology of the roots of some Victorian species. Australian

Journal of Botany 8: 38—50.

READ, D.J. 1983. The biology of mycorrhizas in the Ericales. Canadian

Journal of Botany 61: 985—1004.

READ, D.J. 1991. Mycorrhizas in ecosystems. Experientia 47: 376 —391.

REDHEAD, J. F. 1968. Mycorrhizal associations in some Nigerian forest

trees. Transactions of the British Mycological Society 51: 377-387.

REEVES, F.B., WAGNER, D., MOORMAN, T. & KIEL, J. 1979. The

role of endomycorrhizae in revegetation practices in the semi-

arid west. I. A comparison of incidence of mycorrhizae in severely

disturbed vs. natural environments. American Journal of Botany

66: 6-D.

SAMPSON, K. 1939. Olpidium brassicae (Wor.) Dang, and its connec-

tion with Asterocystis radicis De Wildeman. Transactions of the

British Mycological Society 23: 199—205.

SIEGFRIED, W.R. 1981. Trophic structure of some communities of

fynbos birds. In E.J. Moll, Proceedings of a symposium on coastal

lowlands of the western Cape. University of the Western Cape,

Bellville, South Africa.

SMITH, S.E. & BOWEN, G.D. 1979. Soil temperature, mycorrhizal

infection and nodulation of Medicago truncata and Trifolium sub-

terraneum. Soil Biology and Biochemistry 11: 469—473.

TANSLEY, S. 1982. Koppie conservation project. Unpublished report.

South African Nature Foundation.

TESTER, M., SMITH, S.E. & SMITH, F.A. 1987. The phenomenon

of ‘nonmycorrhizal plants’. Canadian Journal of Botany 65:

419-431.

TRAPPE, J.M. 1987. Phylogenetic and ecologic aspects of mycotrophy

in the angiosperms from an evolutionary standpoint. In G.R. Safir,

Ecophysiology ofVA mycorrhizal plants: 5—25. CRC Press, Boca

Raton, Florida.

VAN DER WESTHUIZEN, G.C.A. & EICKER, A. 1987. Some fungal

symbionts of ectotrophic mycorrhizae of pines in South Africa.

South African Forestry Journal 143: 20—24.

WARCUP, J.H. 1980. Ectomycorrhizal associations of Australian

indigenous plants. New Phytologist 85: 531-535.

WARCUP, J.H. & MCGEE, P. 1983. The mycorrhizal associations of

some Australian Asteraceae. New Phytologist 95: 667—672.

WITKOWSKI, E.T.F. & MITCHELL, D.T. 1987. Variations in soil

phosphorus in the Fynbos Biome, South Africa. Journal of

Ecology 75: 1159-1171.

%

Bothalia 23,1 (1993)

APPENDIX -A preliminary list of the mycorrhizal status of plants occurring in the Cape Floristic Region

* introduced species.

No. = number of specimens examined.

Loc. = locality and dominant vegetation: B = Bainskloof, Mesic Mountain Fynbos; H = Hercules Pillar, West Coast Renosterveld; J = Jonkers-

hoek, Mesic Mountain Fynbos; K = Kirstenbosch Botanical Garden, in cultivation; M = Melkboschstrand, West Coast Strandveld; N

= Nortier Experimental Farm, Lambert’s Bay, West Coast Strandveld; O = Olifantsbos, Cape of Good Hope Nature Reserve, Mesic Mountain

Fynbos; OK = Orange Kloof, Afromontane Forest; P = Pella, Sand Plain Lowland Fynbos; S = Saasveld, George, Afromontane Forest;

pot = plant growing in pot culture.

Lit. = literature references: A & S = this paper; ABL = A.B. Low (1980); AG = A. Gubb cited by ABL; B, M & A = R. Berliner, D.T.

Mitchell & N. Allsopp (1989); EML = E.M. Laughton (1964); FC = F. Coley cited by ABL; H & M = M.T. Hoffman & D.T. Mitchell

(1986); M & R = D.T. Mitchell & D.J. Read (1987); RKR = R.K. Robinson (1973); S & M = C.J. Straker & D.T. Mitchell (1985).

Myc. = mycorrhizal type: VAM = Vesicular-arbuscular mycorrhizas; ERIC = ericoid mycorrhizas; ENDO = endophytic mycorrhizas of an

undesignated type (see text); ORCH - orchid mycorrhizas; ABS = no mycorrhizas seen in specimens examined; DSH = dark septate hyphae.

Bothalia 23,1 (1993)

97

APPENDIX — A preliminary list of the mycorrhizal status of plants occurring in the Cape Floristic Region (continued)

* introduced species.

No. = number of specimens examined.

Loc. = locality and dominant vegetation: B = Bainskloof, Mesic Mountain Fynbos; H = Hercules Pillar, West Coast Renosterveld; J = Jonkers-

hoek, Mesic Mountain Fynbos; K = Kirstenbosch Botanical Garden, in cultivation; M = Melkboschstrand, West Coast Strandveld; N

= Nortier Experimental Farm, Lambert’s Bay, West Coast Strandveld; O = Olifantsbos, Cape of Good Hope Nature Reserve, Mesic Mountain

Fynbos; OK = Orange Kloof, Afromontane Forest; P = Pella, Sand Plain Lowland Fynbos; S = Saasveld, George, Afromontane Forest;

pot = plant growing in pot culture.

Lit. = literature references: A & S = this paper; ABL = A.B. Low (1980); AG = A. Gubb cited by ABL; B, M & A = R. Berliner, D.T.

Mitchell & N. Allsopp (1989); EML = E.M. Laughton (1964); FC = F. Coley cited by ABL; H & M = M.T. Hoffman & D.T. Mitchell

(1986); M & R = D.T. Mitchell & D.J. Read (1987); RKR = R.K. Robinson (1973); S & M = C.J. Straker & D.T. Mitchell (1985).

Myc. = mycorrhizal type: VAM = Vesicular-arbuscular mycorrhizas; ERIC = ericoid mycorrhizas; ENDO = endophytic mycorrhizas of an

undesignated type (see text); ORCH = orchid mycorrhizas; ABS = no mycorrhizas seen in specimens examined; DSH - dark septate hyphae.

98

Bothalia 23,1 (1993)

APPENDIX— A preliminary list of the mycorrhizal status of plants occurring in the Cape Floristic Region (continued)

* introduced species.

No. = number of specimens examined.

Loc. = locality and dominant vegetation: B = Bainskloof, Mesic Mountain Fynbos; H = Hercules Pillar, West Coast Renosterveld; J = Jonkers-

hoek, Mesic Mountain Fynbos; K = Kirstenbosch Botanical Garden, in cultivation; M = Melkboschstrand, West Coast Strandveld; N

= Nortier Experimental Farm, Lambert’s Bay, West Coast Strandveld; O = Olifantsbos, Cape of Good Hope Nature Reserve, Mesic Mountain

Fynbos; OK = Orange Kloof, Afromontane Forest; P = Pella, Sand Plain Lowland Fynbos; S = Saasveld, George, Afromontane Forest;

pot = plant growing in pot culture.

Lit. = literature references: A & S = this paper; ABL = A.B. Low (1980); AG = A. Gubb cited by ABL; B, M & A = R. Berliner, D.T.

Mitchell & N. Allsopp (1989); EML = E.M. Laughton (1964); FC = F. Coley cited by ABL; H & M = M.T. Hoffman & D.T. Mitchell

(1986); M & R = D.T. Mitchell & D.J. Read (1987); RKR = R.K. Robinson (1973); S & M = C.J. Straker & D.T. Mitchell (1985).

Myc. = mycorrhizal type: VAM = Vesicular-arbuscular mycorrhizas; ERIC = ericoid mycorrhizas; ENDO = endophytic mycorrhizas of an

undesignated type (see text); ORCH = orchid mycorrhizas; ABS = no mycorrhizas seen in specimens examined; DSH = dark septate hyphae.

Bothalia 23,1 (1993)

99

APPENDIX— A preliminary list of the mycorrhizal status of plants occurring in the Cape Floristic Region (continued)

* introduced species.

No. = number of specimens examined.

Loc. = locality and dominant vegetation: B = Bainskloof, Mesic Mountain Fynbos; H = Hercules Pillar, West Coast Renosterveld; J = Jonkers-

hoek, Mesic Mountain Fynbos; K = Kirstenbosch Botanical Garden, in cultivation; M = Melkboschstrand, West Coast Strandveld; N

= Nortier Experimental Farm, Lambert's Bay, West Coast Strandveld; O = Olifantsbos, Cape of Good Hope Nature Reserve, Mesic Mountain

Fynbos; OK = Orange Kloof, Afromontane Forest; P = Pella, Sand Plain Lowland Fynbos; S = Saasveld, George, Afromontane Forest;

pot = plant growing in pot culture.

Lit. = literature references: A & S = this paper; ABL = A.B. Low (1980); AG = A. Gubb cited by ABL; B, M & A = R. Berliner, D.T.

Mitchell & N. Allsopp (1989); EML = E.M. Laughton (1964); FC = F. Coley cited by ABL; H & M = M.T. Hoffman & D.T. Mitchell

(1986); M & R = D.T. Mitchell & D.J. Read (1987); RKR = R.K. Robinson (1973); S & M = C.J. Straker & D.T. Mitchell (1985).

Myc. = mycorrhizal type: VAM = Vesicular-arbuscular mycorrhizas; ERIC = ericoid mycorrhizas; ENDO = endophytic mycorrhizas of an

undesignated type (see text); ORCH = orchid mycorrhizas; ABS = no mycorrhizas seen in specimens examined; DSH = dark septate hyphae.

100

Bothalia 23,1 (1993)

APPENDIX— A preliminary list of the mycorrhizal status of plants occurring in the Cape Floristic Region (continued)

* introduced species.

No. = number of specimens examined.

Loc. = locality and dominant vegetation: B = Bainskloof, Mesic Mountain Fynbos; H = Hercules Pillar, West Coast Renosterveld; J = Jonkers-

hoek, Mesic Mountain Fynbos; K = Kirstenbosch Botanical Garden, in cultivation; M = Melkboschstrand, West Coast Strandveld; N

= Nortier Experimental Farm, Lambert’s Bay, West Coast Strandveld; O = Olifantsbos, Cape of Good Hope Nature Reserve, Mesic Mountain

Fynbos; OK = Orange Kloof, Afromontane Forest; P = Pella, Sand Plain Lowland Fynbos; S = Saasveld, George, Afromontane Forest;

pot = plant growing in pot culture.

Lit. = literature references: A & S = this paper; ABL = A.B. Low (1980); AG = A. Gubb cited by ABL; B, M & A = R. Berliner, D.T.

Mitchell & N. Allsopp (1989); EML = E.M. Laughton (1964); FC = F. Coley cited by ABL; H & M = M.T. Hoffman & D.T. Mitchell

(1986); M & R = D.T. Mitchell & D.J. Read (1987); RKR = R.K. Robinson (1973); S & M = C.J. Straker & D.T. Mitchell (1985).

Myc. = mycorrhizal type: VAM = Vesicular-arbuscular mycorrhizas; ERIC = ericoid mycorrhizas; ENDO = endophytic mycorrhizas of an

undesignated type (see text); ORCH = orchid mycorrhizas; ABS = no mycorrhizas seen in specimens examined; DSH = dark septate hyphae.

Bothalia 23,1 (1993)

101

APPENDIX — A preliminary list of the mycorrhizal status of plants occurring in the Cape Floristic Region (continued)

No. Loc. Lit. Myc. Notes

SCROPHULARIACEAE (corn.)

Nemesia

affinis Benth.

versicolor E. Mey. ex Benth.

barbata (Thunb.) Benth.

Phyllopodium heterophyllum (L. f.) Benth.

Polycarena

capensis (L.) Benth.

cephalophora (Thunb.) Le\yns

Sutera linifolia (Thunb.) Kuntze

Zaluzianskya

divaricata Walp.

villosa (Thunb.) F.W. Schmidt

sp.

SELAGINACEAE

Dischisma capitatum (Thunb.) Croisy

Hebenstretia

dentata L.

repens Jarosz

CAMPANULACEAE

Microcodon

glomeratum A. DC.

hispidulum (Thunb.) Sond.

Roella ciliata L.

Wahlenbergia capensis (L.) A. DC.

LOBELIACEAE

Cyphia digitata (Thunb.) Willd.

Lobelia coronopifolia L.

RUBIACEAE

Anthospermum

aethiopicum L.

sp. 1

Galium tomentosum Thunb.

ASTERACEAE

Arctotheca calendula (L.) Levyns

Arctotis

leptorhiza DC.

sp.

Athanasia trifurcata (L.) L.

Cenia turbinata (L.) Pens.

Chrysanthemoides incana (Burm. f.) T. Norl.

Chrysocoma ciliata L.

Cineraria geifolia (L.) L.

Cotula coronopifolia L.

Didelta spinosa (L. f.) Ait.

Dimorphotheca pluvialis (L.) Moench

Elytropappus

glandulosus Less.

rhinocerotis ( L . f. ) Less.

Eriocephalus

sp. 1

sp. 2

sp. 3

Felicia tenella (L.) Nees

Gazania

ciliaris DC.

sp.

Gymnodiscus capillaris (L. f.) Less.

7 P H A & S VAM

12 PMN A&S, B, M&AVAM

ABS in some, Olpidium, other fungi present

ABS in many at P and M, Olpidium. other fungi

present

Or very slight VAM, other fungi present

Or very slight VAM, Olpidium, other fungi present

Other fungi present

Slight VAM infection

Olpidium, other fungi present

DSH, other fungi present

Olpidium, other fungi present

Slight VAM infection

Olpidium present

Slight VAM infection, fine endophyte present

Fine endophyte

Olpidium, DSH, other fungi present

Needs confirmation

Fine endophyte, Olpidium, other fungi present

Other fungi present

Olpidium, other fungi present

ABS in some, Olpidium, other fungi present

* introduced species.

No. = number of specimens examined.

Loc. = locality and dominant vegetation: B = Bainskloof, Mesic Mountain Fynbos; H = Hercules Pillar, West Coast Renosterveld; J = Jonkers-

hoek, Mesic Mountain Fynbos; K = Kirstenbosch Botanical Garden, in cultivation; M = Melkboschstrand, West Coast Strandveld; N

= Nortier Experimental Farm, Lambert’s Bay, West Coast Strandveld; O = Olifantsbos. Cape of Good Hope Nature Reserve, Mesic Mountain

Fynbos; OK = Orange Kloof, Afromontane Forest; P = Pella, Sand Plain Lowland Fynbos; S = Saasveld, George, Afromontane Forest;

pot = plant growing in pot culture.

Lit. = literature references; A & S = this paper; ABL = A.B. Low (1980); AG = A. Gubb cited by ABL; B. M & A = R. Berliner, D.T.

Mitchell & N. Allsopp (1989); EML = E.M. Laughton (1964); FC = F. Coley cited by ABL; H & M = M.T. Hoffman & D.T. Mitchell

(1986); M & R = D.T. Mitchell & D.J. Read (1987); RKR = R.K. Robinson (1973); S & M = C.J. Straker & D.T. Mitchell (1985).

Myc. = mycorrhizal type: VAM = Vesicular-arbuscular mycorrhizas; ERIC = ericoid mycorrhizas; ENDO = endophytic mycorrhizas of an

undesignated type (see text); ORCH = orchid mycorrhizas; ABS = no mycorrhizas seen in specimens examined; DSH = dark septate hyphae.

102

Bothalia 23,1 (1993)

APPENDIX — A preliminary list of the mycorrhizal status of plants occurring in the Cape Floristic Region (continued)

* introduced species.

No. - number of specimens examined.

Loc. = locality and dominant vegetation: B = Bainskloof, Mesic Mountain Fynbos; H = Hercules Pillar, West Coast Renosterveld; J = Jonkers-

hoek, Mesic Mountain Fynbos; K = Kirstenbosch Botanical Garden, in cultivation; M = Melkboschstrand, West Coast Strandveld; N

= Nortier Experimental Farm, Lambert’s Bay, West Coast Strandveld; O = Olifantsbos, Cape of Good Hope Nature Reserve, Mesic Mountain

Fynbos; OK = Orange Kloof, Afromontane Forest; P = Pella, Sand Plain Lowland Fynbos; S = Saasveld, George, Afromontane Forest;

pot = plant growing in pot culture.

Lit. = literature references: A & S = this paper; ABL = A.B. Low (1980); AG = A. Gubb cited by ABL; B, M & A = R. Berliner, D.T.

Mitchell & N. Allsopp (1989); EML = E.M. Laughton (1964); FC = F. Coley cited by ABL; H & M = M.T. Hoffman & D.T. Mitchell

(1986); M & R = D.T. Mitchell & D.J. Read (1987); RKR = R.K. Robinson (1973); S & M = C.J. Straker & D.T. Mitchell (1985).

Myc. = mycorrhizal type: VAM = Vesicular-arbuscular mycorrhizas; ERIC = ericoid mycorrhizas; ENDO = endophytic mycorrhizas of an

undesignated type (see text); ORCH = orchid mycorrhizas; ABS = no mycorrhizas seen in specimens examined; DSH = dark septate hyphae.

Bothalia 23,1 (1993)

103

APPENDIX— A preliminary list of the mycorrhizal status of plants occurring in the Cape Floristic Region (continued)

* introduced species.

No. = number of specimens examined.

Loc. = locality and dominant vegetation: B = Bainskloof, Mesic Mountain Fynbos; H = Hercules Pillar, West Coast Renosterveld; J = Jonkers-

hoek, Mesic Mountain Fynbos; K = Kirstenbosch Botanical Garden, in cultivation; M = Melkboschstrand, West Coast Strandveld; N

= Nortier Experimental Farm, Lambert’s Bay, West Coast Strandveld; O = Olifantsbos, Cape of Good Hope Nature Reserve, Mesic Mountain

Fynbos; OK = Orange Kloof, Afromontane Forest; P = Pella, Sand Plain Lowland Fynbos; S = Saasveld, George, Afromontane Forest;

pot = plant growing in pot culture.

Lit. = literature references: A & S = this paper; ABL = A.B. Low (1980); AG = A. Gubb cited by ABL; B, M & A = R. Berliner, D.T.

Mitchell & N. Allsopp (1989); EML = E.M. Laughton (1964); FC = F. Coley cited by ABL; H & M = M.T. Hoffman & D.T. Mitchell

(1986); M & R = D.T. Mitchell & D.J. Read (1987); RKR = R.K. Robinson (1973); S & M = C.J. Straker & D.T. Mitchell (1985).

Myc. = mycorrhizal type: VAM = Vesicular-arbuscular mycorrhizas; ERIC = ericoid myconrhizas; ENDO = endophytic mycorrhizas of an

undesignated type (see text); ORCH = orchid mycorrhizas; ABS = no mycorrhizas seen in specimens examined; DSH = dark septate hyphae.

104

Bothalia 23,1 (1993)

APPENDIX— A preliminary list of the mycorrhizal status of plants occurring in the Cape Floristic Region (continued)

* introduced species.

No. = number of specimens examined.

Loc. = locality and dominant vegetation: B = Bainskloof, Mesic Mountain Fynbos; H = Hercules Pillar, West Coast Renosterveld; J = Jonkers-

hoek, Mesic Mountain Fynbos; K = Kirstenbosch Botanical Garden, in cultivation; M = Melkboschstrand, West Coast Strandveld; N

= Nortier Experimental Farm, Lambert's Bay, West Coast Strandveld; O = Olifantsbos, Cape of Good Hope Nature Reserve, Mesic Mountain

Fynbos; OK = Orange Kloof, Afromontane Forest; P = Pella, Sand Plain Lowland Fynbos; S = Saasveld, George, Afromontane Forest;

pot = plant growing in pot culture.

Lit. = literature references: A & S = this paper; ABL = A.B. Low (1980); AG = A. Gubb cited by ABL; B, M & A = R. Berliner, D.T.

Mitchell & N. Allsopp (1989); EML = E.M. Laughton (1964); FC = F. Coley cited by ABL; H & M = M.T. Hoffman & D.T. Mitchell

(1986); M & R = D.T. Mitchell & D.J. Read (1987); RKR = R.K. Robinson (1973); S & M = C.J. Straker & D.T. Mitchell (1985).

Myc. = mycorrhizal type: VAM = Vesicular-arbuscular mycorrhizas; ERIC = ericoid mycorrhizas; ENDO = endophytic mycorrhizas of an

undesignated type (see text); ORCH = orchid mycorrhizas; ABS = no mycorrhizas seen in specimens examined; DSH = dark septate hyphae.

Bothalia 23,1: 105-110 (1993)

Pollen morphology of Curroria , Mondia , Socotranthus and Stomato-

stemma (Periplocaceae)

R.L. VERHOEVEN* and H.J.T. VENTER*

Keywords: Curroria, Mondia, Periplocaceae, pollen morphology, Socotranthus, Stomatostemma

ABSTRACT

The pollen morphology of Curroria Planch., Mondia Skeels, Socotranthus Kuntze and Stomatostemma N.E. Br. was studied.

All the genera are characterized by pollen grains arranged in tetrads. The arrangement of the grains may be rhomboidal,

tetrahedral or decussate. The 4—6 pores present are restricted to the junction area of adjacent grains. The exine is smooth.

Exine structure consists of an outer, homogeneous stratum (tectum) subtended by a granular stratum. The intine is well

developed. The pollen grains of tetrads are connected by wall bridges (cross-wall cohesion). Except for small differences

which may occur between species and genera in pollen size and arrangement of tetrads, the pollen is uniform in morphology.

UITTREKSEL

Die stuifmeelmorfologie van Curroria Planch., Mondia Skeels, Socotranthus Kuntze en Stomatostemma N.E.Br. is bestudeer.

A1 die genera word gekenmerk deur stuifmeelkorrels wat in tetrades gerangskik is. Die rangskikking van die stuifmeelkorrels

kan romboidaal, tetraedries of kruisgewys wees. Vier tot ses poriee kom voor en hulle is beperk tot die aansluitingsgebied

tussen aangrensende stuifmeelkorrels. Die eksien is glad en bestaan uit ’n buitenste homogene stratum (tektum) en granulere

stratum daaronder. Die intien is goed ontwikkel . Die stuifmeelkorrels van tetrades is verbind deur wandbrue (dwarswand-

kohesie). Met die uitsondering van klein verskille tussen spesies en genera in stuifmeelgrootte en rangskikking van tetrades,

stem die stuifmeelkorrels in morfologie ooreen.

INTRODUCTION

Periplocaceae and Asclepiadaceae are two closely allied

families with several features in common. The former was

a subfamily (Periplocoideae) of the latter, but raised to

family status by Schlechter (1924), a concept followed by

Bullock (1957), a well known expert in this group. The

Periplocaceae is related to the Apocynaceae on the one

hand and to the Asclepiadaceae on the other. All three

families have in common a milky latex, flowers with

coronas, fruits composed of paired follicles and seeds with

comas of hairs. The Periplocaceae is distinguished by its

spathulate pollen carriers and pollen grains united in

tetrads. This contrasts with single-grained pollen and

absence of pollen carriers in the Apocynaceae, and pollinia

attached to wishbone-shaped translators in the Ascle-

piadaceae.

Of the 50 genera included in the Periplocaceae, approxi-

mately 20 occur in Africa. Among these Raphionacme

Harv. is the largest with 35 species endemic to Africa

(Verhoeven & Venter 1988) and one endemic to Arabia

(Miller & Biagi 1988). Raphionacme is also the only

herbaceous genus, although a number of its species are

climbers. Most of the other genera are lianous and a small

number are shrubs.

Little information is available on the pollen morphology

of the Periplocaceae. Schill & Jakel (1978) investigated

representatives of the following Periplocaceae in their

study on the pollinaria of the Asclepiadaceae: Crypto-

stegia R. Br., Ectadiopsis Benth., Hemidesmus R. Br.,

Omphalogonus Baill., Parquetina Baill. , Periploca L.,

* Department of Botany and Genetics, University of the Orange Free

State, RO. Box 339, Bloemfontein 9300.

MS. received: 1992-08-21.

Raphionacme, Tacazzea Decne. and Zygostelma Benth.

Data on tetrad size and number of pores for nine collec-

tions of Raphionacme, three species of Periploca, two

species of Tacazzea, and one species of Cryptostegia,

Ectadiopsis, Hemidesmus , Omphalogonus, Parquetina and

Zygostelma are also given. Lebrun et al. (1984) in their

identification of Raphionacme bingeri (A. Chev.) Lebrun

& Stork, give SEM results on seven Raphionacme species.

Verhoeven & Venter (1988) have examined the pollen of

35 Raphionacme species; Verhoeven et al. (1989), five

Tacazzea species and the monotypic genus Petopentia

Bullock; and Venter et al. (1990) three species of Ectadium

E. Mey.

In the present study pollen of Curroria Planch., Mondia

Skeels, Stomatostemma N.E. Br. and Socotranthus Kuntze

was examined with special reference to taxonomy. This

paper thus constitutes part of a comprehensive palynologi-

cal investigation and taxonomic revision of the African taxa

of the Periplocaceae currently being undertaken by the

authors.

Taxonomic aspects and distribution of genera

Curroria comprises five species. C. decidua Planch, is

divided into three subspecies (Bullock 1953). With the

exception of C. decidua subsp. decidua which occurs in

southwestern Africa (Angola, Namibia and South Africa),

all the other Curroria species and subspecies occur in the

desert areas of east and northeast Africa (Tanzania, Kenya,

Somalia, Ethiopia, with outlying stations in the

Hadramawt and Socotra) (Bullock 1953). Curroria species

are robust, erect shrubs or scramblers. The linear to

spathulate leaves are borne in clusters on short shoots and

opposite on long shoots. The petiole is absent or very

short. Flowers are solitary or occur as few-flowered cymes

in the axils of the leaves.

106

Bothalia 23,1 (1993)

Socotranthus is a monotypic genus. S. socotranus (Balf.

f.) Bullock is found on the island of Socotra (Suqutra) east

of the coast of Somalia. This is a large woody shrub of

arid habitats. Its suborbicular leaves are opposite and the

white flowers are grouped together in terminal cymes.

Mondia has two species, M. ecomuta (N.E. Br.) Bullock

and M. whitei (Hook, f.) Skeels. Both species are lianas

of moist forest in tropical and subtropical regions of

Africa. They both bear large, cordate leaves and many-

flowered axillary cymes. The flowers are showy, ranging

from yellowish to reddish in colour.

Stomatostemma also comprises two species, 5. mon-

teiroae (Oliv.) N.E. Br. and S. pendulina Venter & D.V.

Field. The first species is a climber with ovate leaves and

large cymes of whitish-purple flowers, whereas the latter

is a virgate shrub with linear leaves and large cymes of

white flowers. Both species occur in dry forest.

MATERIAL AND METHODS

Pollen was obtained from herbarium specimens. For

light microscopy (LM) pollen was acetolysed according

to the method of Erdtman (1960), mounted in glycerine

jelly and sealed with paraffin wax. Samples were examined

with a Zeiss Photomicroscope. Measurements of tetrad

size are based on a minimum of 15 tetrads per specimen.

For scanning electron microscopy (SEM), pollen was

acetolysed, air-dried on stubs, coated with gold and

examined with a Jeol Winsem 6400 microscope. For trans-

mission electron microscopy (TEM) fresh material was

used. Pollen carriers were fixed in 3% phosphate-buffered

glutaraldehyde, postfixed in 1% osmium tetroxide, dehy-

drated in ethyl alcohol and embedded in Spurr’s low-

viscosity resin. Sections were cut with a glass knife,

stained with uranyl acetate, followed by lead citrate, and

examined with a Philips 300 electron microscope at 60 kV.

Pollen specimens examined

Curroria brevifolia Balf. f., no material available.

C. decidua subsp. decidua Planch., Warmbad, Namibia, Mar. 1986,

Beukes 22 (BLFU); Mariental, Namibia, Sept.-Mar. 1959, Werdermann

& Oberdieck 2261 (K); Usakos, Namibia, Feb. 1969, Jensen 93 (WIND);

Ruacana Falls, Namibia, De Winter <6 Giess 7105 (WIND).

C. decidua subsp. gillettii (Hutch. & Bruce) Bullock, no material

available.

C. decidua subsp. volubilis (Balf. f.) Bullock, Kishen, Socotra, Aug.

1956, Gwynne 96 (BM).

C. macrophylla A.R. Smith, Abd al Kuri, Somalia, Oct. 1966, Virzo

A27 (K).

C. migiurtina (Chiov.) Bullock, no locality, no date, Collenette 198

(K); Eil, Somalia, Nov. 1986, Lavranos <& Carter et al. 24945 (K).

C. volubilis (Schltr.) Bullock, Yaida, Tanzania, Jan. 1970, Richards

25306 (K); Harrar, Ethiopia, Nov. 1970, De Wilde 7TJ9 (WAG); Nyambiti,

Tanzania, Mar. 1953, Tanner 1274 (BR).

Mondia ecomuta (N.E. Br.) Bullock, Pangani District, Tanzania, May

1950, Faulkner 558 (K); Kilifi District, Kenya, June 1973, Musyoki <&

Hansen 956 (K); Monyondzi, Congo, Nov. 1964, Bouquet 631 (P).

TABLE 1. — Diameter of pollen tetrads (/zm)

Bothalia 23,1 (1993)

107

FIGURE 1. — SEM of tetrads of species. A, Curroria volubilis, Richards 25306 (K), rhomboidal tetrad. B, C, C. decidua subsp. decidua'. B,

tetrahedral tetrad, De Winter & Giess 7105 (WIND); C, decussate tetrad, Beukes 22 (BLFU). D— F, Mondia whitei: D, rhomboidal tetrad.

Venter 9068 (BLFU); E, tetrahedral tetrad, Scheepers 1058 (PRU); F, decussate tetrad. Venter 9068 (BLFU). G— I, Stomatostemma monteiroae,

Pooley 293 (NU); G, rhomboidal tetrad; H, tetrahedral tetrad; I, decussate tetrad. Scale bars = 10 pm.

M. whitei (Hook, f.) Skeels, Duiwelskloof, South Africa, Nov. 1960,

Scheepers 1058 (PRU); Bloemfontein (glasshouse). South Africa, Dec.

1984, Venter 9068 (BLFU).

Socotranthus socotranus (Balf. f.) Bullock, Aola, Socotra, Mar. 1953,

Popov 275 (BM).

Stomatostemma monteiroae (Oliv.) N.E. Br., Josini, South Africa, Nov.

1983, Venter 8988 (BLFU); Umfolozi Game Reserve, South Africa, Dec.

1961, Ward 3921 (PRE); Ndumu Game Reserve, South Africa, Dec. 1968,

Pooley 293 (NU).

S. pendulina Venter & D.V. Field, Namina, Mozambique, July 1962,

Leach & Schelpe 11441 (K).

RESULTS

Pollen morphology

The pollen grains in the four genera are united in tetrads,

with the grains arranged rhomboidally (Figures 1A, D, G;

2A, D, G), tetrahedrally (Figures IB, E, H; 2B, E, H)

or decussately (Figures 1C, F, I; 2C, F, I; 3). The number

of pores of individual grains of the tetrads may vary, but

the morphology and position are the same for all the genera

investigated. The pores are round, oval or irregular and

are restricted to the junction area of adjacent grains. Pores

are sometimes covered with a thin layer of exine material.

Curroria

Pollen grains are arranged rhomboidally (Figures LA;

2A), tetrahedrally (Figures IB; 2B) or decussately (Figures

1C; 2C). In C. volubilis (Schltr.) Bullock the most com-

mon arrangement is rhomboidal, whereas in the other spe-

cies it is tetrahedral and decussate. Size of tetrads varies

from 27-42 x 22—34 pm (tetrahedral) to 34—55 x

24—33 pm (rhomboidal) (Table 1). Individual grains of

tetrads have 4—6 pores.

Mondia

Pollen grains are arranged rhomboidally (Figures

ID; 2D), tetrahedrally (Figures IE; 2E) or decussately

(Figures IF; 2F). Although rhomboidally arranged grains

are present, the most common arrangements are tetra-

hedral and decussate. Size of tetrads varies from 39-49

x 36—45 pm (tetrahedral) to 48—67 x 38-58

(rhomboidal) (Table 1). Individual grains of the tetrads

have 4-6 pores.

108

Bothalia 23,1 (1993)

FIGURE 2. — LM photographs of tetrads of species. A, Curroria volubilis, Richards 25306 (K), rhomboidal tetrad. B, C, C. decidua subsp. decidua,

Beukes 22 (BLFU): B, tetrahedral tetrad; C, decussate tetrad. D, E, Mondia ecomuta, Faulkner 558 (K); D, rhomboidal tetrad; E, tetra-

hedral tetrad. F, M. whitei. Venter 9068 (BLFU), decussate tetrad. G— I, Stomatostemma monteiroae: G, rhomboidal tetrad. Venter 8988

(BLFU); H, tetrahedral tetrad, Pooley 293 (NU); I, decussate tetrad, Pooley 293 (NU). Scale bars = 10 /im.

B

FIGURE 3. — Decussate tetrad of Soco-

tranthus socotranus, Popov 275

(BM). A, SEM; B, LM photo-

graph. Scale bars = 10 ^m.

Bothalia 23,1 (1993)

Socotranthus

Pollen grains are arranged decussately (Figure 3) or

tetrahedrally, very seldom rhomboidally. The decussate

arrangement differs from that observed in other genera,

in that the cells are arranged more parallel to each other

and not perpendicular. Size of tetrads varies from 28—36

x 22—27 /tm (tetrahedral) (Table 1). Individual grains of

the tetrads have 4-6 pores.

Stomatostemma

Pollen grains are arranged rhomboidally (Figures 1G;

2G), tetrahedrally (Figures 1H; 2H) or decussately

(Figures II; 21). The most common arrangements are tetra-

hedral and decussate. Size of tetrads varies from 29—58

x 27—54 fim (tetrahedral) to 39—61 x 28—51 pm (rhom-

boidal) (Table 1). Individual grains of the tetrads have 4—6

pores.

Exine structure

The exine is smooth and covered with a thin electron-

dense layer (Figure 4A, arrow). Exine structure con-

sists of an outer, homogeneous stratum (tectum) subtended

by a granular stratum (Figure 4B). The tectum and

granular stratum have the same electron density but the

granular stratum has an irregular appearance because of

channels which occur throughout it. The two layers are

separated by a discontinuous line of osmiophilic substance.

The fibrillar intine is well developed. The internal walls

have the same structure as the exterior wall. Wall bridges

consisting of intine and granular layer occur between

adjacent grains (Figure 4C). The intine wall bridges

indicate the position of pores in the internal wall of aceto-

lysed pollen grains.

DISCUSSION

Compound pollen grains occur in more than 56 families

of angiosperms (Erdtman 1945; Walker & Doyle 1975;

Knox & McConchie 1986). The cohesion mechanisms in

mature polyads were discussed by Knox & McConchie

(1986). The cohesion of compound pollen occurs by

attachment of the tectum (simple cohesion) or by connec-

ting wall bridges (cross-wall cohesion). In cross-wall

cohesion wall bridges are present in the common wall

between adjacent grains, and these bridges comprise intine

and granular layer in the Periplocaceae.

Pollen tetrads and polyads are common in a number of

families and have been used in systematic treatments to

separate genera and species (Oldfield 1959; Skvarla et al.

1975; Takahashi 1986). In the Mimosoideae extensive use

of tetrads and polyads is made to separate genera (Guinet

1981a, b; Niezgoda et al. 1983). In the Periplocaceae,

Raphionacme can be distinguished palynologically from

the other genera by the 8—16 pores per pollen grain

(Verhoeven & Venter 1988) as against 8— 10 in Baseonema

Schltr. & Rendle and 4—6 in the other genera. The mono-

typic Petopentia natalensis (Schltr.) Bullock can be

distinguished from the other genera by the presence of

linear and T-shaped tetrad arrangements (Verhoeven

et al. 1989). In Tacazzea, T. tomentosa Bruce differs

significantly from the other species by the width of the

rhomboidal tetrads (Verhoeven et al. 1989).

109

FIGURE 4. — A, B, TEM of pollen wall: A, Curroria decidua subsp.

decidua, Beukes 22 (BLFU); B, Mondia whitei, Venter 9068

(BLFU). C, M. whitei, Venter 9068 (BLFU), internal wall. G,

granular stratum; I, intine; T, tectum. Scale bars = 1 /an.

The present study shows that Stomatostemma pendulina

differs in tetrahedral size (32 x 30 /im) from S. motiteiroae

(48 x 47 /an). In Curroria, C. volubilis differs from the

other species in that rhomboidal tetrad arrangement is

predominant. Socotranthus can be identified by the un-

usual decussate arrangement of the tetrad.

110

Bothalia 23,1 (1993)

The exine structure, consisting of a solid stratum

(tectum) subtended by a granular stratum, appears to be

without much variation in the representatives of the Peri-

plocaceae studied thus far. The exine structure shows a

resemblance with the general exine structure in the Apocy-

naceae. In the Apocynaceae the granular stratum is

however, more variable, e.g. consisting of elements of un-

equal size and shape; subtended by a sole; larger granules

towards the base and partly fused to a sole; faintly defined

and delimited stratum with irregular voids containing

traces of osmiophilic material (Nilsson 1986; 1990).

Although small differences may occur, the pollen mor-

phology of the taxa investigated is rather uniform and thus

of little value in the distinction o> the species and genera

investigated. The different, predominantly rhomboidal

arrangement of the pollen tetrads of Curroria volubilis may

indicate that this species should be placed in a different

genus. It is significant that other floral characteristics show

that C. volubilis does not belong in Curroria or to any

of the genera investigated but to a new genus.

ACKNOWLEDGEMENTS

The financial support by the Foundation for Research

Development and the University of the Orange Free State

is gratefully acknowledged. All the herbaria mentioned

are thanked for the kind loan of their specimens.

REFERENCES

BULLOCK, A. A. 1953. Notes on African Asclepiadaceae. III. Kew

Bulletin 9: 360-362.

BULLOCK, A. A. 1957. Notes on African Asclepiadaceae. VIII. Kew

Bulletin 1956, 12: 503.

ERDTMAN, G. 1945. On the occurrence of tetrads and dyads. Svensk

Botanisk Tidskrift 39: 286-297.

ERDTMAN, G. 1960. The acetolysis method: a revised description.

Svensk Botanisk Tidskrift 54: 561—564.

GUINET, P. 1981a. Comparative account of pollen characters in the

Leguminosae. In R.M. Polhill & P.H. Raven, Advances in legume

systematics 2 : 789 —799. Royal Botanic Gardens, Kew.

GUINET, P. 1981b. Mimosoideae: the characters of their pollen grains.

In R.M. Polhill & R. H. Raven, Advances in legume systematics

2: 835 — 857. Royal Botanic Gardens, Kew.

KNOX, R.B. & McCONCHIE, C.A. 1986. Structure and function of

compound pollen. In S. Blackmore & I.K. Ferguson, Pollen and

spores: form and function : 265-282. Academic Press, London.

LEBRUN, J.P., NILSSON, S. & STORK, A.L. 1984. La veritable

identite du Brachystelma bingeri A. Chev. (Asclepiadaceae).

Bulletin du Museum national d’histoire nature lie, Paris 6:

225-231.

MILLER, A.G. & BIAGI, J.A. 1988. Studies in the flora of Arabia XXIII:

five new species from Oman. Notes from the Royal Botanic

Garden Edinburgh 45 : 61—72.

NIEZGODA, C.I., FEUER, S.M. & NEVLING, L.I. 1983. Pollen

ultrastructure of the tribe Ingeae (Mimosoideae: Leguminosae).

American Journal of Botany 70: 650—667.

NILSSON, S. 1986. The significance of pollen morphology in the

Apocynaceae. In S. Blackmore & I.K. Ferguson, Pollen and

spores: form and function : 359-374. Academic Press, London.

NILSSON, S. 1990. Taxonomic and evolutionary significance of pollen

in the Apocynaceae. Plant Systematics and Evolution, Suppl. 5:

91-102.

OLDFIELD, F. 1959. The pollen morphology of some of the west

European Ericales. Pollen et Spores 1: 19-48.

SCHILL, R. & JAKEL, U. 1978. Beitrag zur Kenntnis der Asclepia-

daceen-Pollinarien. Tropische und subtropische Pflanzenwelt 22:

1-122.

SCHLECHTER, R. 1924. Periplocaceae. In E. Rob & C.E. Fries,

Beitrage zur Kenntnis der Flora des Kenia, Mt Aberdare und Mt

Elgon, V. Notizblatt des Botanischen Gartens und Museums zu

Berlin-Dahlem 9: 23—24.

SKVARLA, J.J., RAVEN, P.H. & PRAGLOWSKI, I. 1975. The evolution

of pollen tetrads in Onagraceae. American Journal of Botany 62:

6-35.

TAKAHASHI, H. 1986. Pollen polyads and their variation in Chimaphila

(Pyrolaceae). Grana 25: 161—169.

VERHOEVEN, R.L. & VENTER, H.J.T. 1988. Pollen morphology of

Raphionacme (Periplocaceae). South African Journal of Botany

54: 123-132.

VERHOEVEN, R.L., VENTER, HJ.T. & KOTZE, I.D.S. 1989. Pollen

morphology of Petopentia and Tacazzea (Periplocaceae). South

African Journal of Botany 55 : 207—214.

VENTER, H.J.T. , KOTZE, J.D.S. & VERHOEVEN, R.L. 1990. A

taxonomic revision of Ectadium (Periplocaceae). South African

Journal of Botany 56: 113—124.

WALKER, J.W. & DOYLE, J.A. 1975. The bases of angiosperm

phylogeny: palynology. Annals of the Missouri Botanical Garden

62: 664-723.

Bothalia 23,1: 111-116 (1993)

Dynamics of the forest vegetation of the Umtiza Nature Reserve, East

London

J.J. MIDGLEY* and P.N. GOBETZ**

Keywords: dynamics, eastern Cape, forests, grain, size-class distribution

ABSTRACT

The forest community at the Umtiza Nature Reserve near East London was surveyed using 24 plots (0.04 ha) in which

all woody stems >0.5 m tall were enumerated. Based on a classification using numbers of stems of canopy species, it was

assumed that basically only one forest community was sampled. Further multivariate analyses suggest that this forest is

fine-grained. Sample plots were similarly placed in ordination space irrespective of whether woody species occurrence was

used as importance value or if species occurrence per size class was used separately [seedlings (0.5 — 1.0 m), saplings (1—5

m) or canopy individuals (>5 m)]. An analysis of size-class distributions of the most common canopy species indicated

that the majority of species exhibited inverse J-shaped size-class distributions. This is the expected pattern for a fine-grained

forest. In these measures of dynamics, this forest is not fundamentally different to the more temperate Afromontane forests.

UITTRJEKSEL

’n Opname is gemaak van die woudgemeenskap in die Umtiza- natuurreservaat naby Oos-Londen; alle houtagtige stamme

>0.5 m hoog is in aanmerking geneem. Gebaseer op ’n klassifikasie waarin aantal stamme van blaredakspesies gebruik

word, is daar aangeneem dat basies slegs een woudgemeenskap ingesluit is. Verdere veelvoudige variantanalises dui daarop

dat hierdie woud fyn gegrein is. Monsterpersele is eenders in ordinasieruimte geplaas ongeag of voorkoms van houtagtige

spesies as belangrikheidswaarde gebruik is en of voorkoms van spesies per grootteklas afsonderlik gebruik is [saailinge (0.5— 1.0

m), jong borne (1—5 m) of blaredak-individue (>5 m)]. Ontleding van grootteklas-verspreidings van die mees algemene

blaredakspesies het getoon dat die meeste spesies omgekeerde J-vormige grootteklas-verspreidings vertoon het. Dit is die

verwagte patroon vir 'n fyngrein-woud. In hierdie metings van dinamika, verskil hierdie woud nie fundamenteel van die

meer gematigde Afromontaanse woude nie.

INTRODUCTION

In brief, forest dynamics is the complex product of inter-

actions between disturbance regime (e.g. type of disturb-

ance, turnover rate), life histories of constituent species

along a shade-tolerant to shade-intolerant continuum and

particulars of the regeneration arena (e.g. regeneration bot-

tlenecks due to biotic or abiotic events) . As such the study

of dynamics needs several inputs. In this preliminary study

we concentrated on aspects of the grain and life history

components of dynamics. By life histories we mean

whether a species is relatively shade-tolerant ‘climax’ or

shade-intolerant ‘pioneer’ and this we have inferred from

size-class distributions.

By grain we mean ‘the mosaic of structural phases’

(Whitmore 1989); or the spatial patterns of seedlings/

saplings and canopy individuals of the different species.

Coarse-grained forests tend to have at least some (recently

disturbed) areas which are dominated by shade-intolerant

species. When plotted in ordination space, coarse-grained

forests should show clear separation of plots according to

whether pioneers or climax species are present, especially

when their sizes are considered. In fine-grained forests

most species can regenerate close to adults and therefore

there are no great differences in species composition or

size-class distributions amongst plots. In other words

* Division of Forest Science and Technology, Jonkershoek FRC, Private

Bag X5011, Stellenbosch 7600.

** Cape Provincial Administration, Umtiza Nature Reserve, PO. Box

5185, Greenfields 5208— deceased 11 February 1992.

MS. received: 1991-12-09.

the successional process occurs on a small spatial scale

in fine-grained forests.

Recent work has shown that in many different forests

at any one time gaps occupy about 1—2% of the area

(Barden 1989; Connell 1989). In forests not subject to

catastrophes these values suggest that stems in the small

size classes of shade-intolerant species should be rare in

comparison to the large bank of advance regeneration of

shade-tolerant species. Thus the size-class distributions

of shade-intolerant species should be flatter than those of

shade-tolerant species. A forest comprised of many species

with relatively flat size-class distributions should thus be

a relatively coarse-grained forest.

Very little has been published on the dynamics of South

African forests. At the moment there is therefore no model

to predict the grain of a South African forest either for

a given environment (e.g. climate and soils) or for forest

structure (e.g. whether there is a preponderance of large

or small-sized individuals). Midgley et al. (1990) indicated

that the southern Cape forests [part of White’s (1983)

Afromontane Forest type] were extremely fine-grained.

At the scale of 0.04 ha plots, most species were able

to recruit continuously and most canopy species were

shade-tolerant. This was interpreted as being due to the

unproductive environment (cool climate, poor soils) and

conservative disturbance regime (few large gaps) which

has favoured shade-tolerant species and restrained both

woody and herbaceous shade-intolerant species.

In contrast to the southern Cape forests, our working

hypothesis for tropical and subtropical gap-phase forests.

112

Bothalia 23,1 (1993)

such as are found at our study site, is that they can be

expected to have a greater component of shade-intolerant

species, of both herbaceous and woody types. This is due

to their situation in a more productive environment (higher

temperatures, summer rainfall) which increases the

opportunities for regeneration of fast-growing pioneer

species. Such forests should therefore be more coarse-

grained due to intermittent recruitment of shade-intolerant

canopy dominants leading to spatially segregated size

classes. Thus an ordination or classification of sample plots

using the occurrence of a species in the seedling, sapling

or canopy size classes, should reveal spatially distinct

groups in a coarse-grained forest. In a fine-grained forest,

where species are able to regenerate close to their own

adults, data of species occurrence by size class should have

little influence on classifications or ordinations.

Our survey was done in a coastal forest of the eastern

Cape. According to Acock's (1988) classification these

forests appear to fall between the Coastal Tropical Forest

Type (Veld Type No. 1) and Valley Bushveld (Veld Type

No. 23). White (1983) mapped these forests as the

Tongaland-Pondoland Forest type and most recently they

have been mapped as Dune Thicket (Lubke, Tinley &

Cowling 1988). Information on forest dynamics in the

eastern Cape is conspicuous by its absence in the over-

view of vegetation in the eastern Cape by Lubke, Tinley

& Cowling (1988) and in Everard (1987).

Our objectives were to use information from plots to:

1, briefly describe the vegetation; 2, compare ordinations

of plots using information on presence/absence of potential

canopy species in the seedling, sapling or canopy layers,

to infer grain; and 3, analyse the size-class distributions

of the important species to infer their life histories.

STUDY AREA

The study area is situated in the Umtiza Nature Reserve

(33°02'S; 27°47'E) which is located on a northeast-facing

slope of the Buffalo River Valley, about 10 kilometres from

East London. Thicket/forest vegetation covers about 550

ha of the reserve. One of the aims of this reserve is to afford

high conservation status to two forest tree species: Umtiza

listeriana Sim, belonging to a monotypic genus of the

Fabaceae, and Buxus macowanii Oliv. , the box-wood much

exploited in the eastern Cape in the past.

Maximum altitude in the reserve is 180 m and many

streams dissect the area giving it a variety of aspects. The

geology of the area has been mapped as Beaufort Group

(Rust 1988) and the soils are mapped as weakly developed

soils on rock with black to brown clays and clay loams

(Hartmann 1988).

At East London, annual precipitation is 919.2 mm which

falls mainly in summer. Mean maximum and minimum

temperatures are 22.7°C and 14.0°C respectively. East

London is extremely windy (mean wind speed of 4.7 ms-1)

as opposed to Cape Town (4.0 ms1) and George

(2.5 ms1). (Climate information from Weather Bureau,

Pretoria.)

METHODS

Sampling

The broad vegetation communities of the Umtiza Nature

Reserve were mapped from aerial photographs and essen-

FIGURE 1. —The location of the 24 sample plots at the Umtiza Nature Reserve in the eastern Cape. The shaded-in areas represent Acacia karroo

grassland and the clear areas forest.

Bothalia 23,1 (1993)

113

TABLE 1.— A species list, based on 24 sample plots, of the forest

vegetation of the Umtiza Nature Reserve

tially two types were found to occur; an Acacia kar-

roo! grassland and a thicket/fores t type. A grid was placed

over a 1:10 000 map of the reserve and 24 sites were select-

ed at random in the forest communities (Figure 1). At each

site a square plot of 20 x 20 m was laid out and all woo-

dy plants >0.5 m tall were enumerated. These plots were

demarcated permanently for further studies on distur-

bance, mortality and growth. At each sample plot each

stem was allocated to one of three size classes: seedlings

(0.5— 1.0 m in height), saplings ( < 5 m in height) and cano-

py (>5 m). Stem diameter at breast height (dbh) was

measured for stems > 5 m. Identifications were made in

the Albany Herbarium, Grahamstown. Nomenclature fol-

lows Gibbs Russell etal. (1987). The forest canopy ranged

from 5— 8 m in height with an occasional emergent up to

15 m. At each plot, slope and aspect were noted and two

soil samples were taken for standard chemical analyses

by the Agricultural Research Institute at Dohne.

Analysis

Initially we classified the forest vegetation using the

multivariate package TWINSPAN (Hill 1979a), with

numbers of stems as the importance values and the default

settings of the package. In total about 60 species occurred

in the canopy (>5 m in height; Table 1) in our 24 plots.

By using species occurrence in seedling, sapling or canopy

levels as separate species, we could increase the apparent

number of ‘species’ to about 150 and the apparent number

of ‘plots’ to 72.

In the second part of this study we used multivariate

techniques to represent grain or succession in space.

Multivariate techniques have long been used for discerning

successional trends, e.g. Fox (1990) used community

trajectories in multivariate space to study mammal

succession over time. We made comparisons between

classifications and ordinations of plots but using the two

data sets (i.e. 60 spp. versus 150 ‘spp.’ and 24 versus 72

‘plots’). The eigenvalues produced by an ordination/classi-

fication give an indication of the variance explained

by an axis [low eigenvalue ( < 0.5) = poor separation of

samples]. The breadth of an ordination axis gives an

indication of the homogeneity amongst samples, with a

value of 400 (4 s.d.) indicating that species occurring at

one end of an axis are almost completely absent at the

other end (Hill 1979b). At this stage there are no rigorous

tests for comparing ordinations, although some progress

has been made (Grossman, Nickerson & Freeman 1991).

We merely visually inspected the package outputs for

114

Bothalia 23,1 (1993)

differences in eigenvalues and breadth of axes, for the

various classifications/ordinations we performed.

We also calculated the position of centroids for ordi-

nations of plots using presence/absence of canopy species

in seedlings, saplings and canopy size classes separately.

A meaningful successional trajectory would be indicated

by significant directional trend in ordination space as

defined by information from each size class.

Our expectations were that in a coarse-grained forest,

significantly different eigenvalues or breadth of axes would

result when data from each size class were used separately,

rather than when information about the canopy layer only,

was used. We expected centroids to be widely separated

in ordination space when species by size-class informa-

tion was incorporated.

For the second part of the study, the DECORANA

(DCA-option)/TWINSPAN packages (Hill 1979a, b) were

again used. Presence/absence (due to the great discrepan-

cies in numbers of stems amongst size classes) was used

as the importance value, and again we used the default

settings of the package.

RESULTS

Environmental data

Soil colours recorded included very dark brown, dark

red-brown, dark brown and brown-black, and soil texture

was sandy clay loam. The soils also had a relatively high

pH and calcium content (Table 2). Slopes were mostly gen-

tle and aspect was predominantly between 90° and 270°.

Vegetation description

Very little phytosociological data have been published

on the eastern Cape forests and thickets. Furthermore,

vegetation patterns are complex (see Acocks 1988). For

this reason we have appended an annotated species list

(Table 1). The maximum number of stems observed per

0.04 ha sample for the canopy size class was 74, for

seedlings it was 454 and for saplings 25. The maximum

number of seedlings per sample plot was high for the

following species: Ptaeroxylon obliquum (421), Buxus

macowanii (189) and Ilex mitis (256). Species richness per

0.04 ha of woody plants ranged from 13—36.

The separation of the forest into types, based on abun-

dance of canopy individuals, was weak (eigenvector =

TABLE 2. — Environmental information pertaining to the study area. For

slope and aspect classes, numbers represent numbers of plots.

Values in brackets indicate the variance

TABLE 3. —Sample plots belonging to first two groups, and associated

eigenvalues, produced by TW1NSPAN using presence/absence

data for canopy species (60 spp.) and canopy species in three

size classes separately (about 150 spp.)

0.376). Furthermore, axis 1 was only a maximum of 2.5

s.d. units broad and axis 2 only 2.3 s.d. units. This

probably reflects the overlapping distribution of the

common species and many rare species (Table 1). Down-

weighting for rare species made a negligible difference

in eigenvalues. We have taken this weak separation of types

and small spread in ordination space of the forest to

indicate that only one vegetation community was present.

Most of the canopy species are well represented in the

seedling and sapling (Table 1) size classes. Essentially

the forest is dominated by Ptaeroxylon obliquum, Harpe-

phyllum caff rum, Olea woodiana, Chaetacme aristata,

Umtiza listeriana and Buxus macowanii (see Table 1).

Vegetation dynamics

The groupings of samples produced at the first division

using 60 species (i.e. only canopy species) versus 150

species (i.e. canopy species per size class separately),

whilst having different orderings, were identical (Table 3)

and both had similar and small eigenvalues (0.26 compared

to 0.29). This indicates that species by size-class data do

not produce a different classification or a different sepa-

ration of samples.

The ordination using 72 ‘plots’ and the 60 canopy species

shows considerable overlap of the three size-class groups

(Figure 2A, B, C). This is clearly depicted by the similarity

of position of each centroid and their relatively close

proximity to the origin. There is no suggestion of orderly

directional trend of movement of centroids. This could

indicate that there is no consistent difference in succes-

sional status amongst plots. The spread along the first

ordination axis is narrow (about 3 s.d.) and the eigen-

vector was also small (0.279).

Size-class distribution

Most of the common canopy species have typical inverse

J-shaped size-class distributions (Table 4). To this group

could be added many of the less common species we

encountered but which had similar size-class distributions.

This includes species such as Maytenus heterophylla,

M. peduncularis, Cussonia spicata, Euclea natalensis,

Brachylaena elliptica and Olea capensis subsp. capensis.

DISCUSSION

We interpret the multivariate analysis to indicate that

this subtropical forest is fine-grained. This forest is there-

Bothalia 23,1 (1993

115

FIGURE 2. — Ordination diagrams using woody species in size classes: A, canopy; B, seedling; C, sapling. All 72 plots were ordinated together

but represented separately for clarity. The dots indicate the centroids. In A, samples 22 and 23 are superimposed.

TABLE 4. — Numbers of individuals per size class of the most common canopy species encountered in the Umtiza Nature Reserve

fore fairly similar to the more temperate southern Cape

forests (Midgley et al. 1990). Both have an abundance of

species with inverse J-shaped size-class distributions and

co-occurrence of different size classes within a species.

These results indicate that large-scale disturbances are not

important in determining community composition of both

these forest types.

From a conservation perspective, because the shifting

mosaic of structural phases (sensu Whitmore 1989) is

fine-grained (or stationary) at Umtiza, relatively small

patches of forest may be viable because most species can

regenerate on a small spatial scale. Obviously, other factors

need to be taken into account (edge effects, visits by

dispersers) to complete the picture. We predict that the

disturbance regime at Umtiza will be conservative (prepon-

derance of small gaps) and that advance regeneration will

inherit most gaps.

Finally, it is clear from this survey that Umtiza listeriana

and Buxus macowanii, the two species of special conser-

vation importance, are both abundant and apparently

regenerating adequately. They appear to be well conserved

in the Umtiza Reserve.

ACKNOWLEDGEMENTS

We thank D. MacDevette for initiating the project. Dept.

Water Affairs and Forestry for funding and D. Everard,

H. van Daalen and especially C. Geldenhuys and anony-

mous referees for comments on the manuscript and

Dominique du Toit for doing much of the sampling.

REFERENCES

ACOCKS, J.RH. 1988. Veld types of South Africa, 3rd edn. Memoirs

of the Botanical Survey of South Africa No. 57.

BARDEN, L.S. 1989. Repeatability in forest gap research studies in Great

Smoky Mountains. Ecology 70: 558, 559.

CONNELL, J.H. 1989. Some processes affecting the species composition

in forest gaps. Ecology 70: 560—562.

EVERARD, D.E. 1987. A classification of the subtropical thicket in the

eastern Cape, based on syntaxonomic and structural attributes.

South African Journal of Botany 53: 329—340.

FOX, B.J. 1990. Changes in the structure of mammal communities over

successional time scales. Oikos 59: 321—329.

GIBBS RUSSELL, G.E., WELMAN, W.G., RETIEF, E., IMMEL-

MAN, K.L., GERMISHUIZEN, G., PIENAAR, B.J., VAN

WYK, M., NICHOLAS, A., DE WET, C., MOGFORD, J.C.

& MULVENNA, J. 1987. List of species of southern African

plants, part 2, edn 2. Memoirs of the Botanical Survey of South

Africa No. 56.

GROSSMAN, G.D., NICKERSON. D. & FREEMAN, M.C. 1991.

Principal component analysis of assemblage structure data: utility

of tests based on eigenvalues. Ecology 72 : 341—347.

HARTMANN, M.O. 1988. The soils of the eastern Cape. In M.N. Bruton

& G.W. Gess, Towards an environmental plan for the eastern

Cape : 37—43. Rhodes University, Grahamstown.

HILL, M.O. 1979a. TWINSPAN—a fortran program for arranging

multivariate data in an ordered two-way table by classification

of the individuals and the attributes. Cornell University, Ithaca,

New York.

HILL, M.O. 1979b. DECORAN A — a fortran program for detrended

correspondence analysis and reciprocal averaging. Cornell

University, Ithaca, New York.

LUBKE, R.A., TINLEY, K.L. & COWLING, R.M. 1988. Vegetation

of the eastern Cape. In M.N. Bruton & F.W. Gess, Towards

an environmental plan for the eastern Cape: 68—87. Rhodes

University, Grahamstown.

MIDGLEY, J.J., SEYDACK, A., REYNELL, D. & MCKELLY, D.

1990. Fine-grain pattern of southern Cape plateau forests. Journal

of Vegetation Science 1: 539—546.

Bothalia 23,1 (1993)

RUST, l.C. 1988. The geology of the eastern Cape. In M.N. Bruton & WHITE, F. 1983. The vegetation of Africa. Unesco.

F.W. Gess, Towards an environmental plan for the eastern Cape: WHITMORE, T.C. 1989. Canopy gaps and the major groups of forest

12—23. Rhodes University, Grahamstown. trees. Ecology 70 : 536 - 538.

Bothalia 23,1: 117-127 (1993)

The vegetation of the northeastern Orange Free State, South Africa:

physical environment and plant communities of the Ea land type

H.C. ECKHARDT*, N. VAN ROOYEN* and G.J. BREDENKAMP*

Keywords: Braun-Blanquet method, classification, geology, Grassland Biome, land types, soils

ABSTRACT

The research was carried out in the Ea land type of the northeastern Orange Free State, with the objective of reclassifying

and refining Acocks’s veld types. TWINSPAN classification results were further refined by Braun-Blanquet procedures. The

100 releves distributed over the Ea land type resulted in the recognition of four major vegetation types which may be divided

into nine plant communities. The communities were hierarchically classified, described and ecologically interpreted.

DECORANA ordination was used to determine vegetation/environmental gradients and relationships.

UITTREKSEL

Navorsing is gedoen op die Ea-landtipe in die noordoostelike Oranje-Vrystaat met die doel om Acocks se veldtipes te

herklassifiseer en te verfyn. Die resultate van die TWINSPAN-klassifikasie is met behulp van Braun-Blanquetprosedures

verder verwerk. Die 100 releves wat oor die hele Ea-landtipe versprei is, het vier hoofplantegroeitipes opgelewer wat in

nege plantgemeenskappe onderverdeel kan word. Die gemeenskappe is hierargies geklassifiseer, beskryf en ekologies geln-

terpreteer. Plantegroei- en omgewingsgradiente is met behulp van DECORANA-ordening bepaal.

CONTENTS

Introduction 117

Study area 117

Geology 118

Physiography 119

Soils 119

Climate 120

Methods 120

Description of communities 121

1. Artemisia afra-Rhus dentata shrubland of the

slopes 121

1.1 Hyperthelia dissoluta— Eragrostis curvula shrub-

land of steep slopes 121

1.2 Hyparrhenia hirta—Diospyros lycioides shrub-

land of moderate slopes 121

2. Themeda triandra—Elionurus muticus grassland

of relatively dry undulating midslopes/plains 124

2.1 Elionurus muticus— Trachypogon spicatus grass-

land of relatively dry, rocky, shallow soils .. 124

Vernonia oligocephala— Trachypogon spica-

tus grassland of relatively dry, rocky, shal-

low soils 124

Harpochloa falx— Trachypogon spicatus grass-

land of relatively moist, rocky, shallow soils . . 124

2.2 Microchloa caffra- Elionurus muticus grassland

of relatively moist soils 124

Tristachya leucothrix— Elionurus muticus varia-

tion 124

Heteropogon contortus— Eragrostis plana varia-

tion 124

3. Themeda triandra— Eragrostis plana transitional

dry/wet grassland 124

4. Eragrostis plana— Eragrostis curvula wet/moist

grassland 124

* Department of Botany, University of Pretoria, Pretoria 0002.

MS. received: 1991-10-29.

4.1 Eragrostis curvula— Setaria sphacelata moist