Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 42,2

Oct. 2012

TECHNICAL PUBLICATIONS OF THE SOUTH AFRICAN NATIONAL

BIODIVERSITY INSTITUTE (SANBI) PRETORIA

Obtainable from SANBI, Private Bag XlOl, Pretoria 0001,

RSA. A catalogue of all available publications will be issued on

request.

BOTHALIA

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 SANBI, 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.

Bothalia Contents: five booklets containing a list of authors

and titles, and an index to taxa and keywords; a) to vols 1-20

(1921-1990); b) to vols 21-25 (1991-1995); to vols 26-30

(1996-2000); d) to vols 31-37 (2001-2007); to vols 38^1

(2008-2011).

FLORA OF SOUTHERN AFRICA (F5A)

A taxonomic treatise on the flora of the Republic of South

Africa, Lesotho, Swaziland, Namibia and Botswana. 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.

Project discontinued.

FSA contributions 1-20 appear in Bothalia'.

1 : Aquifoliaceae. S. Andrews. 1994. Bothalia 24: 163-166.

2: Asphodelaceae/Aloaceae, 1029010 Chortolirion. G.F. Smith.

1995. Bothalia 25: 31-33.

3: Asphodelaceae/Aloaceae, 1028010 Poellnitzia. G.F. Smith

1995. Bothalia 25'. 35, 36.

4: Agavaceae. G.F. Smith & M. Mossmer. 1996. Bothalia 26:

31-35.

5: Buxaceae. H.F. Glen. 1996. Bothalia 26: 37^0.

6: Orchidaceae: Holothrix. K.L. Immehnan. 1996. Bothalia 26:

25^0.

7: Verbenaceae: Vitex. C.L. Bredenkamp & D.J. Botha. 1996.

Bothalia 26'. 141-151.

8: Ceratophyllaceae. C.M. Wilmot-Dear. 1997. Bothalia 27:

125-128.

9: Onagraceae. P. Goldblatt & P.H. Raven. 1997. Bothalia 27:

149-165.

10: Trapaceae. B. Verdcourt. 1998. Bothalia 28: 1 1-14.

1 1 : Zingiberaceae. R.M. Smith. 1998. Bothalia 28: 35-39.

12; Plantaginaceae. H.F. Glen. 1998. Bothalia 2^: 151-157.

13: Ulmaceae. C.M. Wilmot-Dear. 1999. Bothalia 29; 239-247.

14: Cannabaccae. C.M. Wilmot-Dear. 1999. Bothalia 29:

249-252.

15: Piperaceae. K.L. Immehnan. 2000. Bothalia 30; 25-30.

16: Sphenocleaceae. W.G. Welman. 2000. Bothalia 30: 31-33.

17: Casuarinaceae. C.M. Wilmot-Dear. 2000. Bothalia 30:

143-146.

1 8: Salicaceae s. ,str. M. Jordaan. 2005. Bothalia 35: 7-20.

19: Asteraceae: Anthemideae: Eiimorphia. N. Swelankomo.

2011. /io//w//a 41; 277-282.

20: Asteraceae: Anthemideae: Inezia. N. Swelankomo. 2012.

Bothalia 42: 247-250.

STRELITZIA

A series of occasional publications on southern African flora and

vegetation, replacing Memoirs of the Botanical Survey of South

Africa and Annals of Kirstenbosch Botanic Gardens.

• The Memoirs are individual treatises usually of an ecological

nature, but sometimes dealing with taxonomy or economic

botany. Published: Nos 1-63 (many out of print). Discontin-

ued after No. 63.

• The Annals are a series devoted to the publication of mono-

graphs and major works on southern African flora. Published:

Vols 14-19 (earlier vols published as suppl. vols to the Jour-

nal of South African Botany). Discontinued after Vol. 19.

FLOWERING PLANTS OF AFRICA (FPA)

This serial presents colour plates of African plants with

accompanying text. The plates are prepared mainly by the

artists at SANBI. Many botanical artists have contributed to the

series, such as Fay Anderson, Peter Bally, Auriol Batten, Gillian

Condy, Betty Connell, Stella Gower, Rosemary Holcroft,

Kathleen Lansdell, Cythna Letty (over 700 plates), Claire

Linder-Smith and Ellaphie Ward-Hilhorst. The Editor is pleased

to receive living plants of general interest or of economic value

for illustration.

From Vol. 55, twenty plates are published at irregular intervals.

An index to Vols 1^9 is available.

PALAEOFLORA OF SOUTHERN AFRICA

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:

• Molteno Fonnation (Triassic) Vol. 1. Introduction. Dicroidi-

um, 1983, by J.M. & H.M. Anderson.

• Molteno Formation (Triassic) Vol. 2. Gymnosperms (exclud-

ing Dicroidium), 1983, by J.M. & H.M. Anderson.

• Prodromus of South African Megafloras. Devonian to Lower

Cretaceous, 1985, by J.M. & H.M. Anderson. Obtainable

from: A. A. Balkema Marketing, Box 317, Claremont 7735,

RSA.

• Towards Gondwana Alive. Promoting biodiversity and stem-

ming the Sixth Extinction, 1999, by J.M. Anderson (ed.).

• Heyday of the gymnosperms: systematics and biodiversity

of the Late Triassic Molteno fructifications, 2003, by J.M.

Anderson & H.M. Anderson. Strelitzia 15.

• Brief history of the gymnospemis: classification, biodiver-

sity, phytogeography and ecology, 2007, by J.M. Anderson,

H.M. Anderson & C.J. deal. Strelitzia 20.

• Molteno ferns; Late Triassic biodiversity in southern Africa,

2008, by H.M. Anderson & J.M. Anderson. Strelitzia 21.

SANBI BIODIVERSITY SERIES

A series of occasional reports on projects, technologies,

workshops, symposia and other activities initated by or executed

in partnership with SANBI.

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 42,2

Scientific Editor; J.C. Manning

Production Editor: Y. Steenkamp

SANBI

Biodiversity for Life

f

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag XlOl, Pretoria 0001

ISSN 0006 8241

October 2012

Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

M.J.A. Werger

Royal Botanic Gardens, Kew, UK

South African National Biodiversity Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

University of Utrecht, Utrecht, The Netherlands

Acknowledgements to referees

Archer, Mrs C., South African National Biodiversity Institute, Pretoria, RSA.

Barker, Prof. N.P., Rhodes University, Grahamstown, RSA.

Becker, Dr R.W., South African Council for Natural Scientific Professions, Pretoria, RSA.

Berry, Prof. P.E., University of Michigan, USA.

Boatwright, Dr S., South African National Biodiversity Institute, Cape Town, RSA.

Bridson, Dr D.M., Royal Botanic Gardens, Kew, UK.

Fish, Ms L., South African National Biodiversity Institute, Pretoria, RSA.

Gehrke, Dr B., Johannes Gutenberg-Universitat, Mainz, Germany.

Goldblatt, Prof P., Missouri Botanical Garden, St Louis, USA.

Henderson, Ms L., ARC-Plant Protection Research Institute, Pretoria, RSA.

Ivey, Mr P., South African National Biodiversity Institute, Cape Town, RSA.

Klaassen, Ms E., National Botanical Research Institute, Windhoek, Namibia.

Le Maitre, Dr D.C., Council for Scientific and Industrial Research, Stellenbosch, RSA.

Leistner, Dr O.A., ex South African National Biodiversity Institute, Pretoria, RSA.

Linder, Prof R, University of Zurich, Zurich, Switzerland.

Magee, Dr A.R., South African National Biodiversity Institute, Cape Town, RSA.

Manning, Dr J.C., South African National Biodiversity Institute, Cape Town, RSA.

Muasya, Dr M., University of Cape Town, Cape Town, RSA.

Oxelman, Prof B., University of Gothenburg, Gothenburg, Sweden.

Proches, Dr S., University of KwaZulu-Natal, Durban, RSA.

Razafimandimbison, Dr S., Royal Swedish Academy of Sciences, Sweden.

Schrire, Dr B., Royal Botanic Gardens, Kew, UK.

Singh, Dr Y., South African National Biodiversity Institute, Durban, RSA.

Snijman, Dr D., South African National Biodiversity Institute, Cape Town, RSA.

Turland, Dr N., Missouri Botanical Garden, St Louis, USA.

Van Oudtshoom, Mr F., Africa Land-Use Training, Modimole, RSA.

Van Wyk, Prof A.E., University of Pretoria, Pretoria, RSA.

Whitehouse, Dr C., Royal Horticultural Society Garden, Wisley, UK.

Date of publication of Bothalia 42,1: 3 May 2012.

CONTENTS

Bothalia 42,2

1. The emerging invasive alien plants of the Drakensberg Alpine Centre, southern Africa. C.

CARBUTT 71

2. Systematics of the southern African genus Ixia (Iridaceae: Crocoideae): 4. Revision of sect.

D/c/?o«e. P. GOLDBLATT and J.C. MANNING 87

3. Systematics of the hypervariable Moraea tripetala complex (Iridaceae: Iridoideae) of the south-

ern African winter rainfall zone. P. GOLDBLATT and J.C. MANNfNG Ill

4. New species and subspecies of Babiana, Hesperantha, and Ixia (Iridaceae: Crocoideae) from

southern Africa; range extensions and morphological and nomenclatural notes on Babiana

and Geissorhiza. P. GOLDBLATT and J.C. MANNING 137

5. A taxonomic revision of the southern African native and naturalized species of Silene L. (Caryo-

phyllaceae). J.C. MANNING and P. GOLDBLATT 147

6. Notes on African plants:

Cyperaceae. Identity and typification of Carex cognate and status of C. drakensbergensis. C.

ARCHER and K. BALKWILL 190

Fabaceae. Ophrestia oblongifolia (Phaseoleae) in southern Africa. A.N. MOTEETEE and B.-

E. VANWYK 187

Hyacinthaceae. Albuca tenuifolia and^. shawii (Omithogaloideae), two distinct species from

South Africa. M. MARTINEZ- AZORIN, M.B. CRESPO and A.P. DOLD 193

Iridaceae. Notes on Moraea subg. Visciramosa (Iridoideae), including the new species Moraea

saldanhensis from the Atlantic coast of South Africa, recognition of M. inconspicua

subsp. namaquensis, and pollination biology in M. rivulicola. P. GOLDBLATT and

J.C. MANNING 210

Iteaceae. Taxonomic and nomenclatural notes on the genus Choristylis. M. JORDAAN .... 196

Nyctaginaceae. Taxonomic status of Commicarpiis fruticosus. M. STRUWIG and S.J.

SIEBERT 201

Poaceae. The taxonomic and conservation status ofAgrostis eriantha wax. planifolia. J.E. VIC-

TOR, A.C. MASHAU and V.J. NGOBENI 202

Rubiaceae. Taxonomic notes on Sericanthe andongensis and a new combination and status in

Sericanthe from Limpopo, South Africa. M. JORDAAN and H.M. STEYN 204

7. Nomenclature and typification of southern African species of Euphorbia. P.V. BRUYNS 217

8. FSA contributions 20: Asteraceae: Anthemideae: Inezia. N. SWELANKOMO 247

9. South African National Biodiversity Institute: publications 1 January 20 1 1 to 3 1 December 2011.

Compiler: Y. STEENKAMP 251

10. Guide for authors to Bothalia 259

New combinations, names, sections, series, species and statuses in Bothalia 42,2 (2012)

Babiana rivulicola Goldblatt & J.C. Manning, sp. nov. 137

Ser. Euanthera Goldblatt & J.C. Manning, ser. nov., 90

Euphorbia gerstneriana Bruyns, nom. nov., 236

Euphorbia radyeri Bruyns, sp. nov., 228

Hesperantha radiata subsp. caricina (Ker Gawl) Goldblatt & J.C. Manning, stat. nov., 139

Itea sect. Choristylis (Harv.) Jordaan, sect. nov. et stat. nov., 198

Ixia altissima Goldblatt & J.C. Manning, sp. nov., 90

Ixia bifolia Goldblatt & J.C. Manning, sp. nov., 97

Ixia confusa (G.J. Lewis) Goldblatt & J.C. Manning, stat. nov., 97

Ixia flagellaris Goldblatt & J.C. Manning, sp. nov, 107

Ixia minor (G.J. Lewis) Goldblatt & J.C. Manning, stat. nov., 100

Ixia rigida Goldblatt & J.C. Manning, sp. nov., 91

Ixia scillaris subsp. latifolia Goldblatt & J.C. Manning, subsp. nov., 105

Ixia scillaris subsp. toximontana Goldblatt & J.C. Manning, subsp. nov., 106

Ixia simulans Goldblatt & J.C. Manning, sp. nov., 107

Ixia tenuis Goldblatt & J.C. Manning, sp. nov., 106

Ixia teretifolia Goldblatt & J.C. Manning, sp. nov., 140

Moraea cuspidata Goldblatt & J.C. Manning, sp. nov., 120

Moraea decipiens Goldblatt & J.C. Manning, sp. nov., 115

Moraea grandis Goldblatt & J.C. Manning, sp. nov., 133

Moraea hainebachiana Goldblatt & J.C. Manning, sp. nov., 127

Moraea helmei Goldblatt & J.C. Manning, sp. nov., 116

Moraea inconspicua subsp. namaquensis Goldblatt & J.C. Manning, subsp. nov., 212

Moraea mutila (C.H.Bergius ex Eckl.) Goldblatt & J.C. Manning, comb, nov., 119

Moraea ogamana Goldblatt & J.C. Manning, sp. nov., 128

Moraea saldanhensis Goldblatt & J.C. Manning, sp. nov., 210

Moraea tripetala subsp. jacquiniana Goldblatt & J.C. Manning, stat. et comb, nov., 126

Moraea tripetala subsp. violacea Goldblatt & J.C. Manning, subsp. nov., 125

Sericanthe andongensis subsp. legatii (Hutch.) Jordaan & H.M.Steyn, stat. nov., 206

Silene aethiopica subsp. longliflora J.C. Manning & Goldblatt, subsp. nov., 181

Silene burchellii subsp. modesta J.C. Mannmg & Goldblatt, subsp. nov., 170

Silene burchellii subsp. multiflora J.C. Mann/ng & Goldblatt, subsp. nov., 172

Silene burchellii subsp. pilosellifolia (Cham. & Schltdl.) J.C. Manning & Goldblatt, stat. nov., 169

Silene crassifolia subsp. primuliflora (Eckl. & Zeyh.) J.C. Manning & Goldblatt, comb, et stat. nov., 175

Silene rigens J.C. Manning & Goldblatt, sp. nov., 164

Silene saldanhensis J.C. TWbwn/ng & Goldblatt, sp. nov., 160

Silene undulata subsp. polyantha J.C. Maww/ng & Goldblatt, subsp. nov., 160

IV

Bothalia42,2: 71-85 (2012)

The emerging invasive alien plants of the Drakensberg Alpine Centre,

southern Africa

C. Carbutt*

Keywords: biological invasions, climate change, disturbance regime, Drakensberg Alpine Centre, early detection, emerging alien plants, expert

opinion, life history traits, prioritisation, selection criteria, swift management interventions

ABSTRACT

An ‘early detection ’-based desktop study has identified 23 taxa as ‘current’ emerging invasive alien plants in the Dra-

kensberg Alpine Centre (DAC) and suggests a further 27 taxa as probable emerging invaders in the future. These 50 species

are predicted to become problematic invasive plants in the DAC because they possess the necessary invasive attributes and

have access to potentially suitable habitat that could result in them becoming major invaders. Most of the ‘current’ emerging

invasive alien plant species of the DAC are of a northern-temperate affinity and belong to the families Fabaceae and Rosaceae

(four taxa each), followed by Boraginaceae and Onagraceae (two taxa each). In terms of functional type (growth form), most

taxa are shrubs (9), followed by herbs (8), tall trees (5), and a single climber. The need to undertake a fieldwork component is

highlighted and a list of potential study sites to sample disturbed habitats is provided. A global change driver such as increased

temperature is predicted to not only result in extirpation of native alpine species, but to also possibly render the environment

more susceptible to alien plant invasions due to enhanced competitive ability and pre-adapted traits. A list of emerging inva-

sive alien plants is essential to bring about swift management interventions to reduce the threat of such biological invasions.

INTRODUCTION

Increased volumes and frequency of trade, travel, and

tourism have resulted in an increased spread of invasive

and potentially invasive species (Simberloff 2001). The

prediction is that such trends are likely to increase in the

short and medium-term future in South Africa (Richard-

son 2001; Le Maitre et al. 2004). One result of increased

trade, travel, and tourism is that plants will be moved

by humans across geographic barriers far beyond their

natural dispersal range. To date. South Africa has been

invaded by many species of non-native plants, many

of which are already well established and have a nega-

tive ecological and economic impact (Wells et al. 1986;

Richardson et al. 1997; Van Wilgen 2004), particularly

on ecosystem services (Van Wilgen et al. 2008, 2011).

The different rates of spread observed in different areas

are attributed to synergistic interactions between the

basic features of the environment, features of the distur-

bance regime, and life history traits (Richardson 2001;

Thuiller et al. 2006).

Mgidi et al. (2004) predicted that more invasive alien

species are likely to reach South Africa in the immedi-

ate future. Although upon arrival, such species are still

in the infancy of their invasion (either only recently

introduced and/or are entering a phase of rapid popula-

tion growth), they pose an even greater threat than some

of the major established invaders because of the large

areas they have the potential to invade and the ‘unknown

factor’ associated with the exponential phase of their

expansion (Hobbs & Humphries 1995; Nel et al. 2004).

Emerging invaders appear to be establishing in areas

already heavily invaded by major (well-established)

invaders, suggesting that due to certain climatic fea-

tures, patterns of human settlement, and/or land-use pat-

terns, certain areas are more susceptible and predisposed

* Scientific Services, Ezemvelo KZN Wildlife, RO. Box 13053, Cas-

cades 3202, South Afnca. e-mail: carbuttc@kznwildlife.com.

MS. received: 2011-07-19.

to invasive plants in general and that major invaders

are also likely to be facilitating invasions of emerging

invader species (Nel et al. 2004). Alien plant monitor-

ing and management programmes, historically reactive

in nature, should therefore not only target well-estab-

lished invaders; the ‘blacklisting’ of emerging invaders

as an early warning system will help identify, prioritise,

and appropriately manage new invasions (Mgidi et al.

2004) so that the predicted trend of increasing inva-

sions is matched with an ever-increasing ability to nul-

lify the emerging threat (Nel et al. 2004; Dickers 2004).

The overall objective should therefore be to proactively

curb the threat of invasive alien plant species by stop-

ping the invasion in its tracks, which will afford signifi-

cant ‘savings’ in terms of minimising biodiversity losses

and minimising overall management costs. Although the

established invasive alien plants in the DAC are reason-

ably well known and their threat to plant biodiversity in

the region recognised (Carbutt & Edwards 2004, 2006),

no study has focussed explicitly on the emerging inva-

sive alien plants of the DAC and the likely threat that

such invaders may pose in the future.

Emerging invasive alien plants of the DAC are here

defined as either (i) those alien plant species recorded

from the DAC in the past 25 years or less that are cur-

rently still in the early stages of invasion (i.e. less than

100 populations with less than 1 000 individuals per

population) and given their specific attributes and poten-

tially suitable habitat, could expand further to become

major invaders in the future (~ ‘current’ emerging inva-

sive aliens); or (ii) those alien plant species naturalised

in parts of South Africa that do not occur within the con-

fines of the DAC, but in all likelihood will in the future

given either their predominantly temperate affinity and/

or their current range within 70 km’s of the DAC (~

‘future’ emerging invasive aliens).

The aims of this study were threefold: (i) outline a

basic methodology to identify the emerging invasive

alien plants of the DAC; (ii) develop a preliminary list

72

Bothalia42,2 (2012)

using the desktop component of the methodology identi-

fied in (i) above; and (iii) discuss the likely threat posed

by these emerging alien invasive plants and the possible

relationship between their spread and emerging global

change drivers.

METHODS

Scope of study

A desktop approach was adopted to rapidly generate a

preliminary list, which is especially helpful if the scope

of study encompasses a large geographical area (i.e.

where field work would prove costly and time consum-

ing and may take years to complete). The best interim

measure is a list to leverage swift management action

that can later be fine-tuned with field work.

The geographical scope of the desktop study is the

DAC, a temperate region with summer rainfall. Mean

annual rainfall varies from ± 640 mm on the more lee-

ward Lesotho side to over 2 000 mm on the main escarp-

ment (Tyson et al. 1976; Van Wyk & Smith 2001). The

mean temperature of the warmest month is less than

22°C, whilst winter temperatures drop to well below

freezing with snow and frost commonplace. Soils of the

DAC above 3 000 m therefore have frigid or cryic tem-

perature regimes with mean annual temperatures ranging

from 0°C to 8°C (Schmitz & Rooyani 1987). The varied

climate is partly responsible for the 1 1 vegetation types

(five grassland types, five shrubland types, and one for-

est type) occurring in the DAC (see Mucina & Ruther-

ford 2006).

The initial proposed field work component focuses on

the South African portion of the DAC although future

studies should include representative areas of Lesotho. It

is important to note that the term ‘DAC’ is based on cli-

matological and not floristic grounds (Van Wyk & Smith

2001). The delineated area from 1 800 m a.s.l. to the

highest point at 3 482 m a.s.l., encompasses three topo-

ecological zones, namely montane (± 1 300 m to ± 1 900

m), sub-alpine (> 1 900 m to ± 2 800 m) and alpine (>

2 800 m to 3 482 m) zones. The DAC is therefore pre-

dominantly sub-alpine and alpine in nature, with only the

upper limit of the montane zone falling marginally within

the DAC. Therefore many of the emerging invasive alien

plants being discovered in the montane foothills of the

DAC are here listed as ‘future’ emerging aliens as they

currently do not technically occur within the DAC, but in

all likelihood will do so in the future once they breach

the DAC’s climatic envelope.

All sites proposed for the first phase of fieldwork are

dispersed along the length of the Free State, KwaZulu-

Natal and Eastern Cape Drakensberg (Figure 1; Table 1).

These sites are well representative of the eastern DAC,

and take into account a range of disturbance regimes

and environmental heterogeneity related to altitude

(lowest limit to higher altitudes); rainfall (low- to high-

altitude gradients as well as north-to-south latitudinal

and aspects gradients); temperature (low- to high-alti-

tude gradients as well as north-to-south latitudinal and

aspects gradients; see Figure 1) and many of the 11 veg-

etation types occurring within the DAC.

This study only concerns itself with alien plants intro-

duced from areas outside the borders of South Africa

(even though certain species native to South Africa, yet

FIGURE 1 . — Study sites in the eastern DAC proposed for sampling potential emerging invasive plants in disturbed habitats. This figure should also

be interpreted in conjunction with Table I .

Bothalia 42,2 (2012)

73

TABLE 1. — Examples of physically disturbed sites which may facilitate the invasion of certain alien plant species in the DAC. Many such physi-

cally disturbed sites are also sites of nutrient enrichment (e.g. cattle kraals, heavily grazed or overstocked sites, and rural settlements without

on-site or waterborne sanitation).

naturally absent from the DAC, may be viewed as ‘alien’

if introduced into the DAC). A classic example is the

‘Ermelo’ ecotype of Eragrostis curvula (Poaceae) from

the Highveld. Furthermore, certain native species may

even become ‘weedy’ within their native range. Again

such species were excluded [e.g. Artemisia afra, Chryso-

coma ciliata, and Stoebe vulgaris (all of the Asteraceae

family)]. The study also focuses predominantly on ter-

restrial invasive plants, the only exceptions being the

aquatic herbs Glyceria maxima (Poaceae) and Nastur-

tium officinale (Brassicaceae). Plant nomenclature fol-

lows Germishuizen & Meyer (2003).

The floristic and threats analysis was based only on

the ‘current’ emerging invasive alien species identified

in this study.

Two-tiered ‘top-down ’ and ‘bottom-up ’ approach:

introduction

Many alien plant programmes lack objective protocols

for prioritising invasive species and areas based on likely

future dimensions of spread (Rouget et al. 2004) and reli-

able methods of predicting invasion potential are hard

to come by (Nel et al. 2004). Criteria using the impact

scores of Parker et al. (1999) are problematic because

they are essentially qualitative, lacking information relat-

ing to abundance and rates of spread (Le Maitre et al.

2004). Emerging invaders are also not necessarily those

most obvious. For example, species that have the great-

est available habitat and potential to increase in distribu-

tion are sometimes not identified by experts as impor-

tant invaders (Robertson et al. 2004). Many are initially

innocuous and restricted within their introduced range

(Simberloff 2001). This may be because the dynamics

of range expansion and population growth of an invasive

alien plant typically include a time lag between its arrival

in a new habitat and the start of widespread invasion

(Simberloff 2001). Richardson (2001) cited many exam-

ples of invaders not showing invasive tendencies for as

long as ± 50 to 150 years after introduction.

‘Top-down' approach

To avoid ‘reinventing the wheel’, this study drew

in part from the broad-based desktop review of South

Africa’s emerging invaders from natural and semi-natu-

ral habitats (Mgidi et al. 2004, 2007; Nel et al. 2004).

After a screening process, confidence was placed in

a final subset of 28 invaders (out of a possible total of

454) because they had been scrutinised a number of

times by a number of authors. During this process, Nel

et al. (2004) applied the expert scoring of four criteria

strongly associated with factors that predict the poten-

tial invasiveness of plant species (‘impact’, ‘weediness’,

‘bio-control status’ and ‘weedy relatives’) to 454 emerg-

74

Bothalia 42,2 (2012)

ing alien invaders listed in the Southern African Plant

Invaders Atlas (SAPIA) database (Henderson 1998,

2007). In so doing the 454 emerging alien invaders were

reduced to 115, and by further filtering were reduced

to 84 according to estimates of their potential habitat

(‘habitat that can potentially be invaded’) and current

propagule pool size (Nel et al. 2004). Further filtering

by Mgidi et al. (2004, 2007) then reduced the number

to 28 during an exercise identifying the areas in South

Africa with the greatest likelihood of being invaded. The

28 species were also used by Le Maitre et al. (2004) to

assess their potential impacts on the biodiversity, water

resources and productivity of natural rangelands (bush-

veld, grasslands, and shrublands) in South Africa.

The list of 28 species was then scrutinised further for

the DAC context only. The final sub-set amounted to

23 taxa based on expert opinion by the author [personal

field observations, published literature, and specimens

lodged in the Natal University Herbarium (NU) of the

University of KwaZulu-Natal], and distribution records

from Henderson (2001) and the SAPIA database (Hen-

derson 2007). Many of the 23 taxa recognised as ‘cur-

rent’ emerging alien species in the DAC are regarded as

major invaders in other parts of South Africa and there-

fore were not part of Mgidi et al. (2004) and Nel et al.

(2004), as these studies focussed solely on emerging

invasive alien plants.

Although not the primary focus of this study, a further

27 taxa have been tentatively listed as ‘future’ emerging

invasive alien plants in the DAC. These species were

selected on the premise that the next invaders to occupy

the DAC in any meaningful way are most likely those

of a temperate affinity and are currently located within

70 kms of the DAC’s lower altitudinal boundary and

therefore stand the greatest chance of breaching the

DAC’s climatic envelope, or are so poorly known from

few localities that their ‘emerging’ status in South Africa

is yet to be investigated in any detail. These 27 ‘future’

emerging taxa should form the basis of future studies

lest they become forgotten and their future potential eco-

logical threat overlooked.

‘Bottom-up 'approach

Land cover monitoring studies have shown that large

tracts of South Africa’s natural ecosystems are already

transformed (Fairbanks et al. 2000) and the extent and

rate of land transformation will probably increase with

time (Macdonald 1989; Tainton et al. 1989; D. Jewitt

pers. comm.). This trend is in line with other regions of

the world (Dale et al. 1994; Sala et al. 2000). Known for

its high plant species richness and high levels of plant

endemism, the DAC (~ Eastern Mountain hot-spot) is

also characterised by high levels of man-induced habi-

tat transformation and is therefore recognised as one of

southern Africa’s eight biodiversity ‘hot-spots’ (Cowling

& Hilton-Taylor 1994, 1997).

Given the disturbance factor associated with the study

area, the approach takes both disturbed and undisturbed

habitats into account because invasive alien plants are

able to dominate all stages of succession; early (~ sup-

pression) and late (~ tolerance) successional strategies

are contingent upon the specific competitive strategy

employed and can shift in invaded ecosystems over time

(see MacDougall & Turkington 2004). Furthermore, dis-

turbed habitats such as mountain pass roads can extend

the distribution of alien plants beyond reasonable alti-

tudinal expectations (Kalwij et al. 2008). A number of

sites have been proposed for the fieldwork component

(Figure 1; Table 1). These sites may help to identify

additional emerging invasive species undetected in the

‘top-down’ approach, the focus of which is natural and

semi-natural environments.

More specifically, the ‘bottom-up’ approach should

therefore, based on the premise that disturbance is often

a critical prerequisite for the invasion of certain alien

plants, (i) identify major disturbance nodes (sites) as

well as major disturbance types in the DAC (Table 1);

(ii) ensure that the suite of disturbance nodes takes all

major types of disturbance, environmental heterogene-

ity, and land tenure of the DAC into account; (iii) docu-

ment all (potentially invasive) alien plant species at each

designated site; and (iv) compare the field list with the

SAPIA database, keeping in mind that, if not already

data-based, the field species could be an unrecognised

emerging invader. Due to the marked environmental gra-

dients that traverse the length of the DAC from north-

to-south (and east-to-west), certain alien invasive plants

may not be present across all sites (e.g. those of the

drier, colder Eastern Cape Drakensberg vs. those of the

warmer, wetter northern KwaZulu-Natal Drakensberg).

RESULTS

This study has identified 23 ‘current’ emerging inva-

sive alien plant taxa, represented by 15 families, which

pose the most immediate threat to the DAC (Table 2).

The families Fabaceae and Rosaceae contribute most of

the emerging invasive alien species (four taxa each), fol-

lowed by Boraginaceae and Onagraceae (two taxa each).

All other representative families contribute only a single

taxon each (Table 2). In terms of functional type (growth

form), most taxa are shrubs (9), followed by herbs (8),

tall trees (5), and a single climber (Table 2). Of the 23

‘current’ emerging invasive alien plants assessed, some

78% are of a northern temperate affinity, and 22% of

a tropical affinity (Table 3). Interestingly, none of the

assessed taxa were of a southern temperate affinity

{sensu Henderson 2006). All tall trees and almost all

shrubs are of a northern temperate origin, whereas the

herbs are equally representative of northern temperate

and tropical origins.

This study also highlights a further 27 species as

possible ‘future’ emerging alien invasive plants, as the

potential for more recently detected species to invade

into the DAC should not be underestimated (Table 4).

For example, Ruhus phoenicolasius (wineberry), locally

naturalised in the KwaZulu-Natal Midlands (Stirton

1981) may pose a severe problem in the future given the

invasiveness of other Rubus species such as R. cuneifo-

lius in mesic high-altitude grasslands (O’Connor 2005).

Although R. phoenicolasius currently does not occur in

the DAC, it is regarded as a ‘future’ emerging invasive

alien plant given its temperate affinity and proximity to

the DAC (the closest population being ± 70 km’s away).

Bothalia 42,2 (2012)

75

TABLE 2. — The 23 ‘current’ emerging alien plant invaders of the DAC. Taxa are arranged alphabetically.

DISCUSSION

The emerging invasive alien plants of the DAC

Conservatively speaking, at least 170 alien

angiosperm species (or ± 6% of the DAC’s angiosperm

flora) have invaded the DAC (Carbutt & Edwards 2004).

Poaceae and Asteraceae contribute the most established

invaders (Carbutt & Edwards 2004), a reflection of the

general success of these two families in the DAC, and

together with the legume family, Fabaceae, account for

the majority of plant invaders worldwide (Richardson

2001). This study adds a further 23 ‘current’ emerging

invasive alien species and recognises a further 27 taxa

as potential ‘future’ invaders of the DAC. The families

Fabaceae and Rosaceae that together contributed most

of the emerging invasive taxa, also feature as promi-

nent contributors of temperate-affiliated alien invasive

species in southern Africa (see Henderson 2006). Of

the families mentioned above, only the Fabaceae fea-

tures prominently in the native angiosperm flora of the

DAC (Carbutt & Edwards 2004). A possible reason for

the high proportion of alien invasive species of a north-

ern temperate affinity is the long history of cultivating

garden ornamentals from Europe and to a lesser extent

Asia (i.e. many gardens in the foothills of the DAC have

provided the perfect temperate environment for cultivat-

ing ornamental alien plants). More broadly speaking, the

pattern for southern Africa is an equal representation of

taxa from both temperate and tropical origins (Hender-

son 2006). An interesting trend to monitor is the poten-

tial increase in ‘future’ emerging invasive alien plants of

tropical affinity (as a ‘barometer’ of change), given the

global change predictions regarding warming and dis-

placement of native taxa.

The northern and eastern boundaries of the DAC are

estimated to have on average more than 10 alien invad-

ers per quarter-degree grid square, corresponding to

areas with the highest levels of transformation, rainfall,

and population density (Nel et al. 2004). Rouget et al.

(2004) have predicted that most species of emerging

invaders in the DAC will be conflned to the sheltered

conflnes of lower altitudes (montane foothills) below

the escarpment, particularly in the northern KwaZulu-

Natal Drakensberg (warmer and wetter?), the Eastern

Cape and Free State Drakensberg (highly transformed

through agriculture?), and the warmer, sheltered val-

leys of the Lesotho Maloti Mountains. Their absence

from the alpine summit (Lesotho plateau), particularly at

higher altitudes, is attributed to frequent frosts and low

mean temperatures of the coldest month (Rouget et al.

2004). However, although ecosystem invasibility gener-

ally decreases with altitude as fewer alien plants are able

to invade high altitude habitats due to the harsh climatic

conditions (Keeley et al. 2003; Arevalo et al. 2005), a

study by Kalwij et al. (2008) on the distribution of alien

plants along Sani Pass has shown that mountain roads

(particularly verges) are able to increase the altitudi-

nal limit at which an alien plant is invasive due to the

facilitation of a greater propagule pressure, a composite

measure encompassing inter alia the effects of anthro-

pogenically-induced soil disturbance, increased water

runoff, and vehicular traffic (by introducing and spread-

ing propagules). The outcome is the alien species’ ability

to overcome invasion limiting barriers and hence spread

76

Bothalia 42,2 (2012)

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Seeds are also spread by run-off from waterways. Plants may also

regenerate from root and crown fragments left after disturbance.

Taxon (and additional refer- Native range Biogeographical affin- Known range in DAC Potential range in DAC Threat to DAC (consequences of invasion, including target

cnees) ity (per Henderson habitat / predicted niche equivalent and invasive/competi-

2006) tive attributes)

80

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81

to elevations either higher than expected or previously

recorded (Kal^vij et al. 2008).

Which approach is least fallible?

Previous studies have shown that the prioritisation

of invasive species using a ranking system of criteria is

subjective and fallible (see Nel et al. 2004; Rouget et al.

2004) because there are no objective criteria determin-

ing when a score is sufficient to qualify a species for

high-priority management action (Nel et al. 2004). Com-

parisons are also difficult between species that occupy

a wide range of different habitats with varying levels of

disturbance and impact. Robertson et al. (2003) reported

difficulty in ranking priority species requiring manage-

ment action at a local scale, compared to more wide-

spread species (perhaps also less abundant across their

range) requiring intervention over large areas. Rankings

given to species should therefore be viewed as approxi-

mate, rather than absolute (Thorp & Lynch 2000). For

many of these reasons, the two-tiered methodology

using both well-scrutinised scoring systems for invaders

of natural and semi-natural ecosystems and a proposed

fieldwork component to determine the emerging invad-

ers of disturbed habitats has been proposed.

An alternative approach is a predictive one that

makes use of Climate Envelope Models (CEMs). This

approach, however, is also potentially fallible as pre-

dicting the spread of invasive plants is not a perfect sci-

ence because predictions are often subject to numerous

uncertainties (Schneider & Root 2001). A major flaw in

using CEMs is that the role of climate in controlling dis-

tributions is not the same for all species, and other fac-

tors such as disturbance regime and biotic interactions

may sometimes override climatic factors (Richardson &

Bond 1991; Hulme 2003; Le Maitre et al. 2004; Rouget

et al. 2004), particularly when recent introductions are

not in equilibrium with their environment because their

geographic ranges may still be expanding from ‘refu-

gia’ into larger ranges (Rouget et al. 2004). Further-

more, CEMs assume that the current distribution of spe-

cies provides a good indication of their potential range

and the process of averaging climate suitability values

assumes that the mean values represent the location

where the species occurs. This assumption is likely to

be flawed in areas of complex topography (Rouget et al.

2004). CEMs therefore appear to give the best correla-

tions with invasive plant species distributions only at a

national scale (see Rouget & Richardson 2003; Rouget

et al. 2004). Even more disconcerting is the lack of con-

gruency between species selected by expert ratings and

those determined by CEMs (Nel et al. 2004).

Despite the drawbacks of each method, the need to

attempt a list is of paramount importance and the value

of the proactive approach in identifying emerging inva-

sive alien plants cannot be overstated even if there are

no generally accepted ways of quantifying when an area

is ‘invaded’ or when a species is ‘invasive’ (Richardson

2001), and despite the blurry line dividing the emerg-

TABLE 4. — The 27 ‘future’ emerging alien plant invaders of the DAC. Taxa are arranged alphabetically.

82

Bothalia 42,2 (2012)

ing and major invaders. Such a list is useful in (i) help-

ing to select species for modelling their rates of spread;

(ii) knowing what species to target and where to focus

management action in the future; and (iii) facilitating

better trouble-shooting methods for managing biologi-

cal invasions (Nel et al. 2004). The benefits may even

be accrued to the management (control) process itself, as

South African researchers have shown that bio-control is

most effective (Dickers & Hill 1999) and control meas-

ures most cost-effective (Hobbs & Humphries 1995;

Dickers 2004) during the early stages of invasion.

Role of disturbance

Disturbance, be it naturally occurring or human-

induced, is a fundamental driver of plant invasions

(Richardson 2001; Simberloff 2001) because it promotes

characteristic patterns of environmental heterogeneity

and regulates ecosystem processes, population dynam-

ics, species interactions, and species diversity by freeing

up limiting resources (Davis & Moritz 2001). Its effect

is thought to be so critical that some authors (e.g. Elton

1958) have maintained a view that undisturbed native

communities are not susceptible to invasions by intro-

duced species. Irrespective of other factors that facilitate

or limit invasions, the susceptibility of communities to

invasion by alien plants increases with increasing dis-

turbance up to a threshold after which the disturbance

then acts as a barrier — intermediate levels of disturbance

are therefore most optimal for invasiveness (Richardson

2001; Woodward 2001). Invasibility resulting from dis-

turbance is also attributed to reduced competition from

resident plants through the reduction in standing ground

cover (Richardson 2001; Woodward 2001).

The role of disturbance in facilitating the expansion

of emerging invasive alien plants has therefore been rec-

ognised and incorporated into the proposed methodol-

ogy of this study. Early successional invaders are con-

fined mostly to post-disturbance environments and are

effective at acquiring resources in an environment of

high resource availability and relatively low competi-

tion (due to traits such as fast growth), and are therefore

inherently better at suppressing other species in areas of

disturbance (~ suppression-based competition; see Mac-

Dougall & Turkington 2004). Fortunately, the fast-grow-

ing invasives that dominate post-disturbance environ-

ments do not appear to be highly problematic in the long

term because they compete poorly in late-successional

assemblages. Rather, in the absence of disturbance (i.e.

where resources are more limiting in natural or semi-

natural environments), the alien flora is able to toler-

ate reduced resource levels under conditions of intense

competition, thus allowing them to dominate in the latter

stages of succession. This strategy is termed tolerance-

based competition (MacDougall & Turkington 2004).

Opportunities for the spread of invasive plants under

climate change

Climate change is predicted to be one of the great-

est drivers of ecological change in the coming century

(Lawler et al. 2009). The DAC is an excellent laboratory

for the monitoring of climate change because moun-

tainous regions are highly sensitive to environmen-

tal change (e.g. Hill 1996; Midgley et al. 2001), espe-

cially a change in temperature. For example, a major

effect of warming is the tendency of species to track

shifting climate and suitable habitat through dispersal

and migration in order to remain within their optimal

growth environment as present-day plant distributions

are determined by their ecological compatibility with

present-day climate. When climatic conditions change,

plants with a specific set of adaptive characteristics

may no longer be suited to the new conditions (Deacon

et al. 1992; Stock et al. 1997). Consequently, species

are predicted to move poleward in latitude and upward

in elevation (Dunne & Harte 2001). A 3°C change in

temperature is equivalent to a move of 250 km of lati-

tude or 500 m of elevation. Alpine species will tend to

migrate upslope when cooler, higher elevations begin to

warm up (Grabherr et al. 1995; Dunne & Harte 2001;

Kdmer 2001). Species already limited to mountaintops

(i.e. already at their critical physiological threshold) will

be at serious risk of local extinction due to the lack of

potentially suitable habitat to migrate to (Dunne & Harte

2001) and because climates are changing more rapidly

than species can adapt (Schneider & Root 2001). A fur-

ther influence of global warming on alpine plant diver-

sity is the lateral migration of species (~ ‘niche filling’ or

‘horizontal reallocation’), with new niches being filled

by species and other niches being abandoned (Gottfried

et al. 1998; Komer 2001). Ultimately, species that are

similarly affected will occupy similar habitats (Van Zin-

deren Bakker & Coetzee 1988; Hill 1996; Midgley et al.

2001).

Global climate change may not only result in the

direct loss of local native species through climatic

incompatibility; native plant communities will also

become increasingly susceptible to invasions by alien

plants. The impact of alien invasive plants, besides

habitat degradation, is the extinction of native species

through the effects of competition, parasitism, disease,

and hybridisation (Baur & Schmidlin 2007). These

invasive plants either (i) arrive pre-adapted because

the ‘new’ local climatic conditions are similar to what

they experience in their native ranges elsewhere in the

world (Macdonald 1992; Dukes & Mooney 1999), or

(ii) because of some form of change, their competitive

ability increases in their invasive environment. Studies

on plant-climate relationships therefore need to consider

both the current selection pressures as well as future

ones, as currently non-adaptive traits may pre-adapt taxa

to future environmental conditions (Stock et al. 1997).

A future selection pressure in the DAC is warmer tem-

perature, which may be significant given that the cli-

matic boundaries in the DAC are well defined and may

determine the basic distribution limits for plants (Carbutt

2004). The warmer temperatures associated with global

climate change may accelerate organic matter decom-

position and nitrogen mineralisation, thereby creating a

nitrogen environment unsuited to taxa that have evolved

in such nitrogen-limited environments where.soil inor-

ganic nitrogen availability is heavily constrained by

cooler temperatures (Carbutt 2004; Carbutt & Edwards

2008; Contosta et al. 2011). Plant communities thriving

in the DAC’s (currently temperature-mediated) nitrogen-

limited soils may be extirpated by future episodes of

significant warming because of their inability to cope

with nitrogen concentrations far beyond their natural

Bothalia 42,2 (2012)

83

tolerance range. Such communities may therefore be

replaced by common nitrophilous ruderal plants (many

of which will be invasive alien plants) that are neither

conservation-worthy nor native to the DAC.

Management action

Control of invasive alien species is a key operational

management function and demands significant financial

resources. In the uKhahlamba Drakensberg Park World

Heritage Site (UDP WHS), which accounts for a sig-

nificant area of the KwaZulu-Natal Drakensberg portion

of the DAC, some R2 million is spent annually to clear

invasive plants. A concern is that no operational funds

are allocated to combat invasive alien plants in the UDP

WHS; rather all funding is derived from State poverty

relief programmes such as ‘Landcare’, ‘Working for

Water’, ‘Working on Fire’, and ‘Working for Wetlands’,

placing the Park at high risk should this funding be ter-

minated for whatever reason (Ezemvelo KZN Wildlife

2005).

The ‘art and science’ of early detection is a pointless

exercise unless it is followed up with control and miti-

gation by the relevant conservation authorities, and the

failure to detect and eliminate emerging aliens will add

to the financial burden of invasive alien plant control.

Whilst it is acknowledged that areas of high biodiversity

value will be under constant threat by invasive alien spe-

cies and therefore the appropriate management action to

mitigate this threat is an ongoing need (Richardson et al.

2005; Thuiller et al. 2008), the smart approach of early

detection of new invasions is essential to assist manage-

ment in the ongoing battle of alien plant threat mitiga-

tion. Failure to address emerging invasive alien plants in

the short term may double the costs of alien plant con-

trol in the long term. The urgent need to act immediately

and invest a relatively small amount to control emerging

populations before they spread into all available habitats

makes good business sense. The benefits of managing

emerging alien plants should be calculated to include the

stimulation of forther job creation in rural communities,

improved delivery of ecosystem goods and services, and

the safeguarding of native biodiversity and ecological

integrity.

CONCLUSION

South Africa has to its credit a long history of alien

plant control and research (Macdonald et al. 1986;

Van Wilgen et al. 2011). For example, the Working for

Water (WfW) Programme of the Department of Water

and Environmental Affairs (DWEA), initiated in 1995,

has been widely lauded both locally (Van Wilgen et al.

1996; Hobbs 2004; Van Wilgen 2004) and internation-

ally (Mark & Dickinson 2008) for its progressive and

proactive approach in eradicating alien plants. Other

significant allies in the war against invasive alien plants

are the Weeds Research Programme of the Agricultural

Research Council’s (ARC) Plant Protection Research

Institute (PPRI) (which includes the SAPIA database),

and the WfW-funded Invasive Alien Plants Early Detec-

tion and Rapid Response (EDRR) Programme of the

South African National Biodiversity Institute (SANBI).

The critical need for alien plant control and research

in the DAC (relating to both major and emerging invad-

ers) is crucial to maintaining the functional ecological

integrity of the DAC. Like in many other alpine and sub-

alpine environments, the DAC is a mountainous catch-

ment area which supplies drinking and irrigation water,

as well as hydroelectric power. All system functions

(including water quality) is inevitably dependent upon

a healthy and intact cover of vegetation and a species-

rich flora, often likened to an ‘insurance policy’ in that a

species-rich environment has a greater chance of weath-

ering the barrage of human threats and natural environ-

mental changes because there are more likely to be indi-

viduals among the many that can withstand a specific

threat (see Komer 2001). For example, a species-rich

environment has a greater chance of combating invasive

plants because the likelihood of an invader encounter-

ing a close competitor is higher than if just a handful

of native plant species were present (Richardson 2001).

Every conceivable effort should ensure that the levels of

established (major) invasive alien plants in the DAC are

contained to an acceptable minimum and any emerging

alien invasive plants are rapidly identified through early

detection and eradicated before infestation levels impact

negatively on the ecological integrity of the DAC.

FUTURE STUDIES

This desktop-based analysis should be expanded

to include the proposed fieldwork component (to both

verify and potentially widen the net of possible emerg-

ing candidates) and the scope of the assessment enlarged

to include representative areas of Lesotho. The value of

a proactive approach, both in terms of identifying the

invasive alien species and the application of swift coun-

ter measures to eliminate emergent populations before

they become well established, carmot be overstated.

ACKNOWLEDGEMENTS

The author wishes to thank I. Rushworth, S. McKean

and V. Mngomezulu of Ezemvelo KZN Wildlife and Dr.

R. Lechmere-Oertel, previously of the Maloti Drakens-

berg Transfrontier Project, for guidance and sourcing

selected literature. Helpful comments and improvements

from three reviewers are gratefully acknowledged.

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Bothalia 42,2: 87-110(2012)

Systematics of the southern African genus/ja‘« (Iridaceae: Crocoideae):

4. Revision of sect. Dichone

P. GOLDBLATT* and J.C. MANNING**

Keywords: Iridaceae, Ixia sect. Dichone, morphology, new species, southern Africa, taxonomy, winter rainfall zone

ABSTRACT

The southern African genus Ixia L. comprises ± 90 species from the winter-rainfall zone of the subcontinent. Ixia sect.

Dichone (Salisb. ex Baker) Goldblatt & J.C. Manning, one of four sections in the genus and currently including 10 species

and three varieties, is distinguished by the following floral characters: lower part of the perianth tube Aliform and tightly

clasping the style; filaments not decurrent; upper part of the perianth tube short to vestigial; style branches involute-tubular

and stigmatic only at the tips; and so-called subdidymous anthers. We review the taxonomy of the section, providing com-

plete descriptions and distribution maps, and a key to the species. /. amethystina Manning & Goldblatt is recognized to be a

later synonym of I. breviiiiba G.J. Lewis. Most collections currently included under that name represent another species, here

described as I. rigida. We recognize five additional species in the section: early summer-blooming I. altissima from the Ce-

darberg; I. bifolia from the Caledon District; 7. flagellaris, a stoloniferous species from the Cedarberg; I. simulans from the

western Langeberg; and I. tenuis from the Piketberg. We also raise to species rank I. micrandra var. confusa and var. minor,

as 7 confusa and 7. minor respectively. Foliar and associated floral variation in the widespread 7. scillaris has led us to rec-

ognize two new subspecies among its northern populations, broad leaved subsp. latifolia and the dwarfed, smaller flowered

subsp. toximontana; subsp. scillaris is restricted to the immediate southwestern Cape, from Darling to Somerset West. Sect.

Dichone now has 1 7 species and two subspecies.

INTRODUCTION

Restricted to the winter-rainfall zone of southern

Africa, the genus Ixia L. comprises some 90 species

divided among the four sections Dichone (Salisb. ex

Baker) Goldblatt & J.C. Manning, Ixia, Hyalis (Baker)

Diels and Morphixia (Ker Gawl.) Pax (Goldblatt &

Manning 2011). Sect. Dichone, the smallest of these,

comprised eight species and three varieties when last

revised by Lewis (1962). In the later account of Ixia

for Flora of southern Africa, De Vos (1999) followed

Lewis’s taxonomy closely but included the new species,

/. collina, described by Goldblatt & Snijman (1985), and

altered the arrangement of species to place those with

specialized, unilateral stamens at the end of the account.

The most recent addition to the section, /. amethystina,

was described by Manning & Goldblatt (2006), but we

have now discovered it to be conspecific with the type

of I. brevituba. Most plants currently referred to I. brevi-

tuba in herbaria represent a separate species, described

here as /. rigida.

Field work conducted since 1999 has resulted in

the discovery of three sets of populations belonging to

sect. Dichone that do not accord with any of the spe-

cies currently recognized. We describe one of them from

the Langeberg and allied to I. scillaris, as I. simulans,

another from the Piketberg as /. tenuis and the third, evi-

dently allied to the I. micrandra complex, as I. bifolia.

■ B.A. Krukoff Curator of African Botany, Missouri Botanical Garden,

R O. Box 299, St. Louis, Missouri 63166, USA. E-mail: peter.gold-

blatt(gmobot.org.

Compton Herbarium, South African National Biodiversity Institute,

Private Bag XI, Claremont 7735, South Africa / Research Centre for

Plant Growth and Development, School of life Sciences, University of

KwaZulu-Natal, Pietermaritzburg, Private Bag XOl, Scottsville 3209,

South Africa. E-mail: J.Manning(gsanbi.org.za.

MS. received: 2012-05-08.

Examination of living plants and herbarium specimens

matching I. micrandra var. confusa and /. micrandra

var. minor provide evidence that both are better regarded

as separate species, which we treat as /. confusa and I.

minor respectively. We identified two further unde-

scribed species from the Cedarberg in herbaria, namely

I. altissima, an early summer blooming species of uncer-

tain affinities and I. flagellaris, allied to 7. scillaris but

the only stoloniferous species in the section.

We provide a complete review of sect. Dichone,

expanding it to include 17 species, with 7. scillaris sub-

divided into three subspecies.

TAXONOMIC HISTORY

As is the case for many Cape plants, the taxonomic

history of sect. Dichone is complex. The genus Dichone

(the meaning of the Greek translation, two tubes, might

refer to the incompletely dehiscent anther thecae) was

proposed without description by R.A. Salisbury in 1812

as "Dichone crispa Laws. Cat.’ for a plant known at the

time as Ixia crispa L.f (1782) but now treated as 7. eru-

bescens Goldblatt. The cryptic reference, ‘Laws. Cat.’,

refers to an unpublished catalogue, perhaps merely a

handwritten list, of plants evidently grown in the garden

of the Scottish nurseryman, Peter Lawson, about which

little appears to be known. C.F Ecklon (1827) likewise

treated Ixia crispa and its immediate allies in a separate

genus but for which he used the name Agretta, also with-

out description. His species and combinations in Agretta

are currently invalid. Ecklon intended to admit five spe-

cies to his genus, one of which (A. stricta) is now 7.

stricta (Eckl. ex Klatt) G.J.Lewis, and another {A. pal-

lideflavens), which is now 7. odorata Ker Gawl. (sect.

Hyalis according to Goldblatt & Manning 2011). Agretta

stricta was referred by Klatt (1882) to his genus Trito-

nixia and the species was only transferred to Ixia (sect.

Bothalia 42,2 (2012)

Dichone) by Lewis in 1 962. We have not identified Eek-

lon’s A. grandiflora.

The name Dichone was validated by J.G. Baker

(1877) as seet. Dichone Salisb. ex Baker of Tritonia, at

which time it included T. scillaris (L.) Baker, T. triner-

vata Baker and T. nndnlata (Burm.f.) Baker (which he

erroneously believed was an earlier synonym of Ixia

crispa). T. undulata is correctly a species of Tritonia and

is currently known by that name (Goldblatt & Manning

2006). Ixia micrandra Baker (sect. Dichone) remained

in Ixia on account of its actinomorphic flowers, although

it has the filiform perianth tube, short anthers and invo-

lute style branches of Dichone. To add to the series of

misunderstandings, Ixia retusa Salisb., described in

1796 by R.A. Salisbury but without a known type, was

considered conspecific with what Ker Gawler (in Sims

1801) called I. scillaris. Later, Ker Gawler (1803) cited

I. retusa under what he called I. polystachia (sic), which

he then considered an earlier name for /. scillaris. The

plate, t. 629 of Curtiss botanical magazine, is not,

however, /. scillaris but is either /. confusa (G.J. Lewis)

Goldblatt & J.C. Manning or /. stricta (Eckl. ex Klatt)

G.J. Lewis. It appears that Ker Gawler believed that I.

retusa was a member of what we now call sect. Dichone.

Plate t. 629 was later cited by Klatt (1882) under his

combination Watsonia retusa, which like its basionym,

I. retusa, is without a type. Both Lewis (1962) and De

Vos (1999) excluded /. retusa from Ixia on the basis that

the protologue is inadequate to identify the species. We

concur.

The four species of sect. Dichone that occur in the

western Karoo remained unknown until the early twen-

tieth century, when collections of the three species Ixia

curvata G.J. Lewis, I. rigida Goldblatt & J.C.Marming,

and I. trifolia G.J.Lewis were made by Rudolf Marloth

in 1920. The first of the western Karoo species to be

recognized, I. trifolia, was described by Lewis (1934),

based on her own collection grown at Kirstenbosch

Botanical Garden. In assigning the species to Ixia, Lewis

evidently did not then realize that 7. trifolia was closely

allied to species of sect. Dichone, which at the time was

included in Tritonia. Although she noted the unusual

anther dehiscence via very narrow slits in /. trifolia, she

regarded the species as belonging in sect. Euixia (i.e.

sect. Ixia) and close to I. ovata (Andrews) Sweet, now /.

abbreviata Houtt.

By 1954 Lewis regarded sect. Dichone as more

closely allied to Ixia than to Tritonia and recommended

its recognition as a separate genus (Lewis 1954: 100)

because of its unique anther dehiscence. Later (Lewis

1962) she formally transferred sect. Dichone to Ixia,

noting that the section was also unusual in the genus

in its involute-tubular style branches and its distinctive

anthers. She described I. brevituba G.J.Lewis and I.

curvata in 1962, assigning Marloth’s collection of what

we here describe as 7. rigida to her 7. brevituba (Lewis

1962), which is typified by a 1929 collection made by

Grant & Theiler.

Time has provided new evidence for Lewis’s conclu-

sion that Dichone is more closely related to Ixia than to

Tritonia. Species examined cytologically have the same

basic chromosome number, x = 10, and karyotype as

other species of Ixia whereas Tritonia has x = 1 1 (Gold-

blatt 1971a; De Vos 1982; Goldblatt & Manning 2011).

Ixia also has derived pollen grains with a single banded

operculum (Goldblatt et al. 1991 and unpublished)

whereas Tritonia has the plesiomorphic pollen grains

of subfamily Crocoideae with a two-banded operculum.

The two genera are clearly closely allied and available

molecular studies (Goldblatt et al. 2006, 2008) indicate

a sister relationship (with moderate bootstrap support,

80%) between the one species each of Ixia (7. latifolia

D.Delaroche) and Tritonia {T. disticha (Klatt) Baker)

that were sequenced. Although Goldblatt & Manning

(1999) initially treated sect. Dichone as one of two sec-

tions of subg. Ixia they later reverted to Lewis’s four-

section classification of Ixia. There have been no molec-

ular studies that shed light on the relationship of Ixia

and Dichone but the unstated assumption implicit in the

current taxonomy is that Dichone is nested within Ixia,

hence its status as a section of that genus.

MATERIALS AND METHODS

Using standard methods of taxonomic investiga-

tion, we examined the holdings of Ixia in herbaria with

significant southern African collections, BOL, K, MO,

NBG, PRE, and SAM (acronyms following Holmgren et

al. 1990). Our herbarium studies were accompanied by

field investigation to determine variation within popula-

tions for some species, and their ecology, especially soil,

aspect, and altitude.

MORPHOLOGY

Morphology, the vegetative morphology of sec.

Dichone corresponds closely to that of other species of

Ixia, with few exceptions. Leaves are plane and form a

basal fan in most species but 7. erubescens has crisped

and undulate leaf margins, and leaves of 7. scillaris are

sometimes undulate, or rarely crisped in a few popula-

tions from the Piketberg, Pakhuis-Biedouw Mtns, and

near Tulbagh. Leaves of 7. minor (= 7. micrandra var.

minor) are terete to subterete or plane whereas those of

7. micrandra are filiform, and both species usually have

two or sometimes three leaves, the uppermost ± entirely

sheathing the lower part of the stem. 7. bifolia, I. trin-

ervata, and usually 7. simulans also have two leaves,

a basal foliage leaf and a second, largely to entirely

sheathing, upper leaf In 7. trinervata the lanceolate foli-

age leaf is distinctive in having three ± equally promi-

nent veins. The lowermost leaf of 7. simulans has one

prominent vein and thickened margins, and the upper

sheathing leaf is inserted above the stem base.

Flowers are produced in spikes typical of Crocoideae.

The floral bracts are typical of Ixia, being membranous

and ± translucent, becoming dry with age. The outer

of each bract pair usually has 3 prominent veins and 3

teeth but is occasionally 1 -toothed, sometimes varying

within a species but consistently so in 7. rigida. Flow-

ers of most species are predominantly pale to deep pink,

sometimes mauve pink, but with a white to pale yellow

throat often edged darker pink to mauve. The perianth is

occasionally white in some individuals or populations

Bothalia 42,2 (2012)

89

of I. scillaris, notably in the Olifants River Valley popu-

lations of subsp. laiifolia, but is purple in 1. brevituba,

the flowers of which have a dark centre and stamens.

Flower orientation ranges from fully upright to half to

fully nodding, thus facing to the side. Flowers of several

species are scented. I. bifolia, I. confusa, I. stricta, and

at least some populations of I. scillaris produce a rose-

like scent, and /. collina and I. rigida are also sweetly

scented (Goldblatt & Snijman 1985).

The perianth tube is usually described as filiform (or

slender throughout) in sect. Dichone but the tepals are in

fact fused for some distance beyond the lower filiform

part of the tube. The length of the upper part of the tube

is often so short as to be conveniently ignored but in Ixia

altissima and I. rigida fully half the length of the tube is

flared for ± 2 and 1 mm respectively and the tube is thus

clearly funnel-shaped. In I. trifolia the flared upper part

of the tube is also developed to a significant degree. The

walls of the slender, lower part of the tube always tightly

clasp the style and the tube contains no nectar.

Although radial floral symmetry is the common, and

by inference ancestral, condition in the group, flowers

of Ixia collina, I. erubescens, I. simulans, I. scillaris, I.

tenuis, and probably also I. flagellaris are weakly zygo-

morphic. The flowers in these species face sideways and

are thus nodding, the tepals held more-or-less vertically,

with the stamens unilateral and the anthers horizon-

tal or drooping. At dehiscence the stamens in the nod-

ding flowers of I. curvata and I. stricta are symmetri-

cally arranged but become unilateral with age, evidently

through gravity (De Vos 1999).

Filaments are usually free and filiform, and inserted

at the mouth of the narrow part of the perianth tube. In

some populations of Ixia trifolia the filaments are united

basally for up to 0.7 mm, or up to one fifth their length.

Anthers are oblong to subrotund and slightly to mark-

edly recurved above the base. Species from the West-

ern Karoo typically have oblong anthers, 4—5 mm long,

with the thecae ± twice or more as long as wide, whereas

those from the southwestern Cape below the Escarpment

are characterized by their derived, short, suborbicular

to broadly oblong anthers, 2-4 mm long and < twice as

long as wide. Several species within this group have uni-

lateral stamens (/. collina, I. erubescens, I. simulans, and

I. scillaris), with the thecae sharply acute at their bases.

The filaments in these species are characteristically

sharply recurved apically such that the thecae are held

almost horizontally. Anthers in sect. Dichone dehisce

from the base via narrow slits that do not open fully, a

unique feature in the genus and family. Dehiscence is

either ± complete to the locule apex or incomplete and

restricted to the lower third or half of the anther, as in

I. scillaris. Anthers have been described as subdidymous

(Lewis 1962), a term meant to imply the vestigial nature

of the anther cormective, which in sect. Dichone is a nar-

row, adaxial strip of tissue that continues from the apex

of the filament to shortly below the anther apex. The

two thecae are closely appressed without the usual well-

developed sterile tissue of the cormective on their adax-

ial surfaces.

The filiform style divides at the mouth of the narrow

part of the perianth tube, thus at or shortly beyond the

filament bases, and the style branches spread outwards,

either ± straight (e.g. I. bifolia, I. micrandra, I. trifolia)

or falcate to recurved. The style branches are involute,

with the margins rolled upward to form an enclosed,

tubular charmel open only at the apex, with the stig-

matic surfaces restricted to the apices, which are often

expanded or slightly bifurcate (e.g. in I. collina and I.

micrandra).

The capsules are, as far as known, typical of Ixia

(Goldblatt & Manning 2011), as are the seeds, which

have a smooth surface and an excluded vascular trace

(Goldblatt et al. 2006; Goldblatt & Manning 2011). Cap-

sules are known for only a few species and we do not

routinely include capsule features in the species descrip-

tions below. Notably, however, the few available cap-

sules of I. scillaris are ± obovoid, possibly a distinguish-

ing taxonomic feature.

INFRASECTIONAL CLASSIFICATION

No classification below sectional rank has until now

been proposed for sect. Dichone. However, with 17

species, it seems useful to us to recognize two species

groups, or series. We include species with radially sym-

metric flowers and either complete or incomplete anther

dehiscence in ser. Euanthera, thus including the Cedar-

berg I. altissima, all western Karoo species, and several

from the southwestern Cape. The anther thecae in ser.

Euanthera dehisce conventionally along narrow slits for

their entire length or almost so and are either ± rounded

or acute and recurved at the base. Species with mainly

zygomorphic flowers and ± unilateral stamens, I. stricta

excepted, are included in ser. Dichone. The anthers in

this group of species are mostly horizontal or pendent,

acute and recurved at the base, and dehisce incompletely

from below. No vegetative specializations consistently

accompany these floral differences. Ser. Dichone, which

is evidently monophyletic, is centred in the southwestern

Cape below the Escarpment, with populations of I. scil-

laris ranging into northern Namaqualand and onto the

Gifberg-Matsikamma Mtn complex. Ser. Euanthera is

comparatively widespread.

SYSTEMATICS

Ixia sect. Dichone (Salisb. ex Baker) Goldblatt &

J.C. Manning in Bothalia 29: 63 (1999). Tritonia sect.

Dichone Salisb. ex Baker: 163 (1877). Tritonia subg.

Dichone (Salisb. ex Baker) Baker: 190 (1892). Trito-

nixia sect. Dichone (Salisb. ex Baker) Klatt: 357 (1882).

Ixia subg. Dichone (Salisb. ex Baker) G.J. Lewis: 150

(1962). Type: I. scillaris L., lectotype here designated

[Lewis: 159 (1962) and Goldblatt & Manning: 63

(1999) listed I. crispa L. (or its synonym 7. erubescens

Goldblatt) as the type but only Tritonia undulata (with

its presumed synonym /. crispa) and I. scillaris were

included by Baker in the section when it was first validly

described].

Dichone Lawson ex Salisb.: 320 (1812), nom. nud.

Agretta Eckl.: 23 (1827), nom. nud.

90

Bothalia 42,2 (2012)

Conn as in Ixia', stolons with a terminal cormlet

produced in I. flagellaris. Leaves as in Ixia but some-

times only two and then lower with expanded blade and

upper largely to entirely sheathing. Spike: outer bracts

with 3(1) main veins acutely 3-toothed, 3-lobed, or

1 -toothed. Flowers pink to magenta (rarely white), tepal

bases white or yellow edged darker pink (blue-mauve

with dark centre in /. brevituba), scented or unscented;

perianth tube usually relatively short, filiform below and

clasping style sometimes fiirmel-shaped with upper part

flared. Stamens symmetrically disposed or unilateral

and reclinate, filaments inserted at apex of narrow part

of tube, filiform or flattened; anthers straight and linear

to oblong or short and suborbicular, with vestigial con-

nective, thecae fully or incompletely dehiscent, then

splitting from base by narrow slits. Style dividing at

mouth of tube opposite or just above base of filaments.

branches filiform-tubular and involute, thus stigmatic

only at apices, straight or falcate-recurved.

Sen Euanthera Goldblatt & J.C. Manning, sen nov.

Flowers upright or nodding. Stamens symmetri-

cally arranged (at least in bud and on opening, some-

times later by gravity unilateral in Ixia curvata); anthers

straight, rounded at base, or recurved and ± acute at

base, dehiscing conventionally by longitudinal slits to

reach apex or almost so. Style branches ± straight or fal-

cate. Type species: /. micrandra Baker.

10 spp., western half of the Cape floral region and

western Karoo.

1. Ixia altissima Goldblatt & J.C. Manning, sp. nov.

TYPE. — Western Cape, 3219 (Wuppertal): Gonna-

Key to species of sect. Dichone

la Plants 0.9- 1.2 m high; perianth tube funnel-shaped with flared upper portion ± 2 mm long and ± as long as filiform lower half; style

dividing opposite middle 1/3 of anthers 1. /, altissima

lb Plants 0.9-1. 5 m high; perianth tube cylindrical almost throughout, flared only in upper 0. 5-1.0 mm if at all; style dividing opposite or

just above bases of filaments:

2a Flowers radially symmetric, upright to half nodding; stamens erect with anthers parallel or diverging:

3a Perianth tube very short, 2. 0-2. 5 mm long; stem with short, stiffly erect lateral branches held close to main axis, each with up to 8

flowers 2.1. rigida

3b Perianth tube 2. 3-6.0 mm long; stem simple or with 1 (-several) suberect to spreading branches:

4a Leaves ± terete or filiform, rarely more than 1 mm wide; corm tunics of fine fibres extending upward in a collar around the stem base:

5a Style branches straight and extending outward below anthers, 3-5 mm long; filaments filiform; outer floral bracts with 3 main

veins and several secondary veins, acutely 3-toothed 9.1. micrandra

5b Style branches falcate, recurved at tips, up to 2 mm long; filaments flattened and wider in middle; outer floral bracts with 3 main

veins but lacking secondary veins, usually shallowly 3-toothed or 3-lobed 10. /. minor

4b Basal and sometimes other leaves linear to sword-shaped or lanceolate, usually at least 1.5 mm wide, blades plane with main vein

and sometimes multiple veins ± equally prominent; corm tunics fine to coarse-textured, with or without fibrous collar around the

stem base:

6a Anthers 4—5 mm long, oblong and > twice as long as wide; spike nodding in bud, becoming erect as flowers open:

7a Anthers yellow; perianth pink, tepals darker at base and yellow in throat; foliage leaves (2.5-)5-12 mm wide; style branches

4—5 mm long 2.1. trifolia

7b Anthers purple; perianth light purple with small dark centre; foliage leaves 1. 5-2.0 mm wide; style branches up to 2 mm long

4. /. brevituba

6b Anthers 1 .5-3.0 mm long, suborbicular to broadly oblong and < twice as long as wide; spike erect in bud:

8a Leaves 2, basal leaf with expanded, ± lanceolate blade, upper leaf entirely sheathing:

9a Basal leaf with 3 ± equally prominent, thickened veins; style branches falcate to recurved (white drying ± white); tepals

12-15 X 6-8 mm when fully open; spike mostly 7-1 2-flowered 6. 1. trinervata

9b Basal leaf with 1 prominent vein (translucent when alive, appearing raised when dry); style branches ascending, ± straight

(pink, often drying purple); tepals remaining slightly cupped, 12-14 x 5-7 mm; spike mostly 4— 7-flowered 7.1. bifolia

8b Leaves > 2, usually 3 to 5, with no visible veins or only main vein prominent:

10a Leaves (3)4 or 5, often drying at anthesis; foliage leaf blades lanceolate, mostly 4—9 mm wide 11./. stricta

10b Leaves (2)3 or 4; foliage leaf blades linear, mostly 1 .5-A.O mm wide 8. /. confusa

2b Flowers usually half to fully nodding, facing to side; stamens unilateral with anthers held horizontally or pendent:

1 la Anthers dehiscing completely by longitudinal slits 5.1. curvata

1 lb Anthers dehiscing by short slits from base to middle, not reaching apex:

12a Foliage leaf usually single, sometimes 2 (uppermost leaf ± entirely sheathing), blade ± linear, 1.0-4. 5 mm wide; flowers usually

4— 8 per spike; perianth tube ± 4.5 mm long 16. 1, simulans

12b Foliage leaves 3-7, blades lanceolate to falcate, mostly 4—20 mm wide, or closely undulate and then only 2^ mm wide; perianth

tube 2. 5^.0 mm long:

1 3a Leaves 2-4(5) mm wide; corm tunics of fine, soft fibres; perianth tube 2-3 mm long:

14a Leaf margins strongly undulate and crisped; anthers ± 1.5 mm long 12. /. erubescens

14b Leaf margins plane; anthers 2-3 mm long:

15a Corms producing slender stolons from base, terminating in a single, large cormlet; anthers ± 3 mm long 15. 1, flagellaris

15b Corms not producing stolons; anthers 2. 0-2. 5 mm long 14. /. tenuis

1 3b Leaves 4—25 mm wide and margins plane or sometimes undulate(-crisped), corm tunics of firm, medium-textured to coarse

fibres; perianth tube 2. 5-5.0 mm long:

16a Plants to 90 cm high, often branched, with branches held at right angles to main axis, becoming suberect distally; filaments

expanded toward apex and becoming as wide as anthers; style branches widely flared at tips 17. /. collina

16b Plants rarely more than 60 cm high, simple or branched, with branches ascending to suberect; filaments ± uniformly filiform,

< 1/2 as wide as anthers; style branches not noticeably flared at tips 13. /. scillaris

Bothalia 42,2 (2012)

91

fontein, seep in sandstone ground among Kniphofia, 900

m, (-CB), 5 Dec. 2005, Pond 303 (NBG, holo.).

Plants 0.9-1. 2 m high, stem surrounded at base by

membranous cataphylls, simple or with 1-2 branches

subtended by translucent, attenuate bracts and prophylls

up to 8 mm long. Corm 15-18 mm diam., mature tunics

not known. Leaves 4, lower 3 ± basal, reaching to ± mid-

dle of stem, blades linear to linear-sword-shaped, (3-)4-

10 mm wide, firm-textured, margins and midrib and

often a pair of secondary veins thickened and hyaline

(at least when dry), uppermost leaf largely sheathing.

Spike erect, elongate, weakly flexuose, 14-20-flowered,

lateral spikes 6-14-flowered; bracts dry, membranous,

translucent, flecked with brown in upper half, outer ± 7

mm long, usually with 3 prominent veins and 3 -toothed,

inner bracts ± as long as outer or slightly shorter,

2-veined and 2-toothed. Flowers rotate, mauve-pink;

perianth tube narrowly funnel-shaped, ± 4 mm long, fili-

form and clasping style below, flared in distal ± 2 mm;

tepals spreading, oblong-ovate, 12-13 x ± 4 nim. Sta-

mens symmetrically disposed, erect; filaments filiform,

inserted just above narrow part of tube, ± 4 mm long,

± white; anthers oblong, ± 3 mm long, erect, dehiscing

fully by longitudinal slits, yellow. Style dividing oppo-

site middle of anthers, branches ± 2 mm long, falcate,

pale yellow, tubular, stigmatic apically, arching outward

between or shortly overtopping anthers. Flowering time\

Nov.-mid-Dec.

Distribution and habitat: Ixia altissima is known

from two sites in the central and southern Cedarberg,

from the Matjies River Nature Reserve and from Gonna-

fontein to the south (Figure 1). Plants grow along stream

banks or in marshy seeps on sandstone derived soil, at

the Gormafontein site among Kniphofia plants. The habi-

tat remains moist at least until December and possibly

later into the summer. We suggest a conservation status

for I. altissima of Rare (R), in light of its evident rarity,

but we see no current threat to the species.

Diagnosis and relationships: Ixia altissima is a sur-

prising plant, sometimes standing over 1 m in height

and flowering late in the season, well into December.

The anthers with vestigial cormective and the invo-

lute style branches, stigmatic only at the tips, indicate

its placement in sect. Dichone but the funnel-shaped

FIGURE 1. — Distribution of Ixia altissima, •; /. rigida, o; L curvata,

▲ ; and I. stricta, A.

perianth tube with the filaments inserted in the middle

of the upper part of the tube is unusual for the section,

most species of which have the tube virtually filiform

throughout with the filaments inserted at the apex of the

tube. The perianth tube, only ± 4 mm long, and the pale

floral bracts are reminiscent of two Roggeveld Escarp-

ment species, I. trifolia and /. rigida, the latter also flow-

ering relatively late in the season, often in November.

These species share relatively unspecialized, longitudi-

nally dehiscent anthers. /. altissima seems most like /.

rigida, also a tall plant, up to 600 mm high and some-

times flowering as late as November. The latter has flow-

ers with a similarly funnel-shaped perianth tube 2-3 mm

long and silvery translucent floral bracts, contrasting

with the slightly longer perianth tube, ± 4 mm long, and

shorter anthers, just 3 mm long, in I. altissima, which is

also unusual in the section in having the style dividing

opposite the middle of the anthers.

Representative specimens

WESTERN CAPE.— 3219 (Wuppertal): Cedarberg,

Matjies River Nature Reserve, stream bank, (-AD), 21

Nov. 1999, Low 5838 (NBG); Cedarberg, Gonnafontein,

seep in sandstone ground among Kniphofia, 900 m, (-

CB), 3 Dec. 2000, Pond 256 (NBG)

2. Ixia rigida Goldblatt & J.C. Manning, sp. nov.

TYPE. — ^Northern Cape, 3220 (Sutherland): ± 38

km north of Sutherland to Middelpos, Farm Geelhoek,

rocky dolerite slope with red clay soil, 3 108 ft [950 m],

(-AB), 7 Sept. 2006, Goldblatt & Porter 12796 (MO,

holo.; NBG, iso.).

Plants (150-)300-600 mm high, stem usually with

2-3 ± erect branches held close to axis, sheathed at base

by firm, chestnut brown cataphylls, combining with old

corm tunics and leaf bases to form collar around base.

Corm 12-18 mm diam., tunics of firm, relatively hard

fibres. Leaves (3-)5-7, lower (2-)3-6 with expanded

blades, upper 1 or 2 leaves sheathing lower 1/3 to mid-

dle of stem, blades sword-shaped to sub-linear, (3-)5-8

mm wide, main vein, margins, and often secondary vein

pairs moderately thickened, hyaline when dry. Spike

weakly flexuose, nodding in bud, mostly 8-15-flowered,

lateral spikes with fewer flowers; bracts membranous,

silver-translucent, ± 5. 5-7.0 mm long, outer with promi-

nent brown or purple central vein and few secondary

veins not reaching upper margin, acute or with 2 obscure

lateral lobes, inner 2-veined and acutely 2-toothed.

Flowers rotate, pale pink, darker at tepal bases and ±

white in throat, sweetly rose-scented; perianth tube fun-

nel-shaped, ± 2.5 mm long, filiform and clasping style

in basal 1 .5 mm or less, widely flared distally for up to 1

mm; tepals spreading, ovate-oblong, ± 12(-15) x 5(-7)

mm, obtuse. Stamens symmetrically arranged, erect; fila-

ments inserted at mouth of narrow part of tube, filiform,

± 1.5(2. 5) mm long, ± white; anthers oblong to linear,

curved back at base, 3^(5) mm long, slightly recurved

basally, thecae subacute at base, dehiseing fully by

longitudinal slits, yellow. Style dividing at base of fila-

ments, branehes ± straight, 2. 0-3. 0(3. 5) mm long, ± fil-

iform-tubular, stigmatic apically, white. Flowering time:

Sept, to mid-Nov. Figure 2.

92

Bothalia 42,2 (2012)

FIGURE 2. — Ixia rigida, Goldhlatt & Porter 12796 (NBG). A, whole

plant; B, outer (left) and inner (right) bract; C, half flower. Scale

bar: 10 mm. Artist: John Manning.

Distribution and habitat: Ixia rigida is restricted to

the Roggeveld, where it is known from the Escarpment

edge on the Farms Fransplaas, Quaggasfontein, Uitkyk,

Houdenbek and a short distance inland northwest of

Sutherland on the Farm Geelhoek (Figure 1). Plants are

restricted to dolerite outcrops and grow in rocky ground

in the heavy red clay characteristic of this habitat. The

first record of the species, made by Rudolf Marloth in

1920 and said to be from the ‘Komsberg flats ’[some

distance south of Sutherland] may be erroneous, for

7. rigida is otherwise unknown south of the town. The

relatively few records show that the species is rare but

plants often flower late, usually after mid-October on the

escarpment edge, when relatively little collecting is done

in the Roggeveld, so the present records may not fully

reflect its range. There appears to be no imminent threat

to its survival but at sites where we have seen the spe-

cies, plants were heavily overgrazed and often do not set

seed as a result. We tentatively suggest a conservation

status of near-threatened, (NT).

Diagnosis and relationships: Ixia rigida is a distinc-

tive species easily recognized by the moderately tall

stem up to 600 mm high with 2-3 upright branches held

close to the axis, and the silvery, translucent floral bracts

subtending very short-tubed flowers, the tube up to 2.5

mm long with the narrow portion up to ±1.5 mm long.

Plants have up to seven leathery leaves with prominently

thickened margins and main and secondary veins. I. rig-

ida was known to Lewis (1962) from only a single spec-

imen, which she associated with 7 brevifolia and De Vos

( 1 999) likewise misunderstood the species, citing speci-

mens of 7 rigida under the name 7 brevifolia.

Although fairly uniform across its range the popula-

tions of Ixia rigida inland of the Roggeveld escarpment

and flowering three to four weeks earlier than those

along the edge of the escarpment, thus in September,

have slightly larger flowers and less fibrotic leaves. Too

little is known about 7 rigida to assess the significance

of this variation. We note, however, that the escarpment

populations mostly have 6 or 7 leaves, typically 5-7 mm

wide, tepals ±12x5 mm, anthers ± 3 mm long, and style

branches ± 2 mm long. The inland populations have 4

or 5 leaves, mostly 7-8 mm wide, tepals ± 14—16 x 6-8

mm, slightly longer filaments, anthers 4(5) mm long, and

style branches ±3.5 mm long. The bracts of the inland

plants also stand out in their darker purple veins.

Representative specimens

NORTHERN CAPE. — 3220 (Sutherland): Sutherland, Bo-Visrivier

road. Farm Fransplaas, stony dolerite flats, (-AB), 19 Oct. 1995, Gold-

blatt & Manning 10370 (MO); Roggeveld Escarpment, Farm Quag-

gasfontein, dolerite ridge near farm buildings, 4 900 ft [± 1 580 m],

(-AB), 1 1 Oct. 2004, Goldblatt & Porter 12667 (MO); Farm Uitkyk,

Naaldegraskop, (-AD), 14 Nov. 1987, Goldblatt & Manning 8645

(MO, NBG, PRE); Houdenhoek, Hoedenbek 4x4 trail, among dol-

erite boulders, 1 387 m, (-AD), 10 Oct. 2008, Clark & Coombs 880

(NBG). Without precise locality and possibly an error: Komsberg flats,

Oct. 1920, Marloth 9782b (PRE).

3. Ixia trifolia G.J. Lewis in Flowering Plants of

South Africa 14: t. 543 (1934); Lewis: 171 (1962); De

Vos: 69 (1999). Type: South Africa, [Western Cape],

Tweedside, cultivated in Cape Town, Sept. 1932, Lewis

s.n. (Nat. Bot. Card. 2706/32 in BOL, lecto., here des-

ignated; BOL!, K, PRE!, SAMI, isolecto.) [Lewis: 171

(1962) cited this number at BOL as the holotype but

there are two sheets, hence our designation of a lecto-

type].

Plants mostly 150-300 mm high, stem usually with

l-2(-3) short branches, or simple, sheathed at base by

short, firm, green cataphylls. Corm mostly 10-18 mm

Bothalia42,2 (2012)

93

diam., tunics of coarse fibres, vertical elements often

becoming claw-like ridges. Leaves 3 or 4(5), lower 2 or

3 with expanded blades, lanceolate to falcate, 1/2-2/3 as

long as stem, (2.5-)5-8(-12) mm wide, central vein and

margins moderately thickened, upper 1(2) leaves sheath-

ing lower 1/3-2/3 of stem, without expanded blade.

Spike weakly flexuose, inclined, drooping in bud, mostly

3-7-flowered; bracts membranous, translucent, 5-6 mm

long, outer with prominent central vein, sometimes with

2 prominent secondary veins, acute or trilobed, inner

slightly shorter than outer, 2-veined and 2-toothed.

Flowers rotate, pale to deep pink (rarely mauve), yel-

low or white edged with darker pink at base of tepals,

unscented; perianth tube funnel-shaped, 3. 5^.0 mm

long, filiform in basal 2. 5-3.0 mm and clasping style,

flared in distal 1 mm, tepals spreading, ovate-elliptic,

obtuse to retuse, 12-15 x 5. 0-6. 5 mm. Stamens sym-

metrically arranged, erect, filaments inserted at mouth of

narrow part of tube, 3-4 mm long, free or united basally

for up to 0.7 mm, filiform, ± white, anthers oblong-lin-

ear, 3-4 mm long, yellow, thecae obtuse at base, dehisc-

ing fully by longitudinal slits. Style dividing between

base and lower 1/3 of filaments, style branches 4—5 mm

long, projecting between filaments, gently arched out-

ward, filiform-involute, stigmatic apically, pale pink.

Flowering time: mid-Aug. to late Sept. Figure 4E, F.

Distribution and habitat: Ixia trifolia has a relatively

wide range, extending along the Roggeveld Escarpment

from the Farm Uitkyk southward through the Klein Rog-

geveld to the Laingsburg Karoo as far west as Tweedside

(Figure 3). Plants grow on stony clay ground in low, kar-

roid scrub or mountain renosterveld. The flowers offer

no nectar and are pollinated by pollen-collecting female

bees. The anthophorine Amegilla spilostoma (Api-

dae) has been captured visiting flowers near Sutherland

(Goldblatt & Manning 2007) and we have also noted

small hopbine beetles consistently visiting flowers at a

population on the Farm Blesberg and clearly actively

transferring pollen from flowers of one plant to those of

another.

Diagnosis and relationships: Ixia trifolia is recog-

nized by the combination of bright pink, rarely mauve

flowers, white to yellow edged with dark pink in the

centre, always held upright on inclined spikes, and two

or three, relatively broad foliage leaves plus one (or two)

more sheathing the stem. The specific epithet trifolia

refers to the three foliage leaves, thus ignoring the pres-

ence of the upper, entirely sheathing leaf Like other spe-

cies of sect. Dichone from the Western Karoo, the style

branches are slender and nearly straight and the anthers

dehisce completely. Unique to this species in the section

is that the filaments may be united basally for up to one

quarter of their length, and the style is exserted up to 1.5

mm from the filiform part of the perianth tube. Ixia trifo-

lia is perhaps most closely allied to I. brevituba, which

has similarly oriented, upright flowers on an inclined

spike but with narrower leaves, up to 1 .5 mm wide and

violet flowers darker in the centre. Both species stand

out in the section in having the spikes drooping in bud.

Ixia trifolia is relatively common across its range but

is poorly collected. The earliest collection of the spe-

cies that we have found was made by Rudolf Marloth

in 1 920 at the top of Verlatekloof south of Sutherland

although the type population was discovered at Tweed-

side near Touws River sometime before 1932. The type

specimens comprise plants cultivated at Kirstenbosch

Botanic Gardens under G.J. Lewis’s name.

Representative specimens

NORTHERN CAPE. — 3220 (Sutherland): Roggeveld Escarpment,

entrance to Noudrif Farm, NW of Sutherland, (-AB), 23 Sept. 1981,

Goldblatt 6344 (MO); Farm Voelfontein, banks of Bo-Visrivier, (-

AB), 11 Oct. 2004, Goldblatt & Porter 12665 (MO); Farm Koorlands-

kloof 70, N of Kruiskloof, (-AB), 26 Sept. 2009, Helme 6202 (NBG);

Farm Hottentotskloof, SW of Sutherland, (-AC), 2 Oct. 1 999, Gold-

blatt & Ndnni 11193 (MO; PRE); top of Verlatekloof, ,Farm Jakkals-

vlei, (-AC), Oct. 1920, Marloth 9646 (PRE); Klein Roggeveld, flats

along stream at Farm Oranjefontein, (-DC), 30 Sept. 2004, Goldblatt

& Porter 12660 (MO).

WESTERN CAPE. — 3220 (Sutherland): Klein Roggeveld, Kruis-

pad, at rnmoff from Matjiesfontein-Sutherland road, (-DC), 19 Sept.

2003, Goldblatt & Porter 12311 (MO). 3320 (Montagu): Memorial,

flats W of the cemetery, (-AB), 9 Sept. 2006, Goldblatt & Porter

12811 (MO).

4. Ixia brevituba GJ.Lewis in Journal of South Afri-

can Botany 28: 170 (1962); De Vos: 70 (1999), in part.

Type: South Africa, [Northern Cape], ‘Quagas [sic] Pass

[probably Ganagga Pass], road between Middelpos and

Ceres’, Sept. 1929, Grant & Theiler 4931 (BOL, holo!).

I. amethystina J.C. Manning & Goldblatt: 139 (2006),

syn. nov. Type: Northern Cape, 3220 (Sutherland): west

of Farm Agterkop, near top of Gannaga Pass, 16 Sept.

1997, Goldblatt & Manning I0745A (NBG, holo.!, MO!,

iso.).

Plants 150-300 mm high, stem either unbranched and

then with scale-like leaves at aerial nodes, or with up to

2 short branches, erect below, inclined above, sheathed

below by submembranous, red-brown cataphylls. Corm

12-20 mm diam., tunics of medium-textured, wiry,

reticulate fibres, extending upward in papery collar

30-70 mm long. Leaves 3 or 4, basal, uppermost com-

pletely sheathing lower 2/3 of stem, blades suberect or

lowermost slightly falcate, 1. 5-2.0 mm wide, reaching

to near top of stem, firm-textured, margins and midrib

hyaline, lightly thickened. Spike inclined, nodding in

bud, crowded, 5-7 -flowered, branches fewer-flowered;

bracts dry-membranous, translucent or flushed purple

above, outer 5-7 mm long, acute or obscurely 3 -den-

tate, inner ± as long as outer or slightly shorter, 2-veined

FIGURE 3. — Distribution of Ixia bifolia. A ; 1. brevituba. A; 1. trifolia,

•; and /. trinervata, o.

94

Bothalia 42,2 (2012)

and 2-toothed, margins connate in lower 1.5 mm around

ovary. Flowers rotate, light purple with small dark pur-

ple stain in centre, lightly scented; perianth tube ± fun-

nel-shaped, ± 2.3 mm long, filiform below and clasp-

ing style, flaring in distal 1 mm; tepals spreading and

slightly concave distally, obovate, ± narrowed below

into a short claw, 12-13 x 7-8 mm. Stamens symmetri-

cally arranged, erect, filaments inserted at apex of tube,

diverging above, 2.5 mm long, blackish purple, anthers

± oblong, 4. 5-5.0 mm long, blackish purple, dehiscing

laterally by narrow slits along entire length. Style divid-

ing at or slightly above base of filaments, ± 2 mm long.

FIGURE 4. — Ixia brevituba,

NBG192794 (NBG). A, flow-

ering plant; B, outer (left) and

inner (right) bract; C, flower;

D, stamens and style branches.

/. trifolia, Helme 1662 (NBG):

E, flower; F, stamens and style

branches. Scale bars: A, C, E,

10 mm; B, 5 mm; D, F, 2 mm.

Artist: John Manning.

branches filiform, involute, stigmatic at apex, purple,

falcate, 2.0-2. 5 mm long. Flowering time: late Sept, to

early Oct. Figure 4A-D.

Distribution and habitat: Ixia brevituba is known

from a small area on the edge of the Roggeveld Escarp-

ment near Ganagga Pass southwest of Middelpos (Fig-

ure 3). The few known collections were made within

a few kilometres of each other on Farms Zoekop and

Agterkop (the type is not localized exactly but is very

likely from the same area). Plants grow in stony, light

clay in renosterveld. The flowers open widely around

Bothalia 42,2 (2012)

95

08:00, even in cool, overcast conditions and remain fully

open until late afternoon. Flowers of the closely related

I. trifolia, like many other species of sect. Dichone, will

not open fully or at all under cool conditions. The only

pollinators recorded are small hopliine beetles, Kabou-

sia axillaris. The form of the flowers is consistent with

this pollination system. In view of its narrow range we

concur with Raimondo et al. ’s (2009) conservation sta-

tus of vulnerable (VU), but we note that the entire range

of 1. brevituba falls within the Tanqua National Park and

future disturbance seems unlikely.

Diagnosis and relationships: distinguished by its pale

purple flowers with small, dark eye, Ixia brevituba has

relatively broad, blackish anthers that dehisce laterally

so that the light-coloured pollen forms a contrasting pale

margin to each anther, and short style branches that are

± as long as the filaments, thus scarcely projecting from

between the filaments. The lovely amethyst-coloured

flowers are borne on inclined spikes so that they face

directly upward in an elegant, arching spray. I. brevituba

falls among a small group of species that are endemic or

near-endemic to the Roggeveld Escarpment and Klein

Roggeveld, including /. curvata, I. rigida, and I. trifo-

lia, all of which share relatively unspecialized, longi-

tudinally dehiscent anthers 3-5 mm long. Among these

species I. brevituba appears to be most closely allied to

I. trifolia on the basis of their unusual, inclined spikes,

and lateral branches that are decurved and nodding when

young. All other species have spikes that are erect from

bud to fhxit, a feature not explicitly mentioned by Lewis

(1962) in the protologue but noted and illustrated by De

Vos (1999). The two species also share a perianth tube

2-3 mm long. I. brevituba differs from 7. trifolia in its

blackish purple anthers and purple perianth tube, which

gives the flowers a small, dark central eye, consistently

narrower leaves 1. 5-2.0 mm wide, and medium-textured

corm tunics drawn into a well-developed neck. I. trifo-

lia, like all other species in sect. Dichone, has yellow

anthers, although the filaments may be pale mauve, and

a pale perianth tube, giving the flowers a well-defined

whitish central eye. The leaves of I. trifolia are broader

than in I. brevituba, (2. 5-)5. 0-12.0 mm wide, and the

tunics are more coarsely fibrous, with the lower fibres

developed into woody claws but not drawn into a neck

above. In addition, the style branches of I. trifolia are

longer than the filaments, 4-5 mm long, and project con-

spicuously between them.

The ranges of Ixia brevituba and I. trifolia comple-

ment one another — only a short distance separates the

two species and we infer that they share a common

ancestor. The differences in their flowers appear to be

related to their pollination biology. I. brevituba has flow-

ers that conform to the hopliine beetle pollination syn-

drome (Goldblatt et al. 1998) and the beetle Kabousia

axillaris has been captured while visiting the flowers.

All individuals of this insect carried pure loads of Ixia-

type pollen on their dorsal thorax and frons (Manning &

Goldblatt 2006). In contrast, Ixia trifolia is pollinated by

solitary female anthophorine bees (Goldblatt & Manning

2007) and possibly also by hopliine beetles.

When described by Lewis (1962), Ixia brevifolia

included a collection of a second species, now 7. rigida,

and the protologue is a composite of the two species.

De Vos (1999) likewise misunderstood 7. brevituba and

later collections that she associated with the name are

also 7. rigida. Following De Vos’s (1999) interpretation

of 7. brevituba. Manning & Goldblatt (2006) described

7. amethystina for plants that we now realize conform

exactly to the type of 7. brevituba. That name now falls

into synonymy.

Representative specimens

NORTHERN CAPE. — 3220 (Sutherland): Farm Zoekop, 3 km S

of entrance along road to Gannaga Pass, (-AA), 26 Sept. 2002, NBG

192794 (NBG); Farm Zoekop, past ruins near edge of escarpment,

(-AA), 24 Sept. 2002, Rosch 154 (NBG); W of farm Agterkop, near

the top of Gannaga Pass, 16 Sept. 1997, Goldblatt & Manning 10745 A

(MO, NBG).

5. Ixia curvata G.J. Lewis in Journal of South African

Botany 28: 171 (1962); De Vos: 70 (1999). Type: South

Africa, [Northern Cape], Agterhantamsberg, Moorde-

naarspoort, 26 miles [± 39 km] NE of Calvinia, 25 Sept.

1952, Lewis 3532 (SAM, holo.!; BOL [SAM 61919]!,

PREl, iso.).

Plants 120-350 mm high, stem simple or with 1 or

2 upright branches, sheathed at base by long, chestnut-

brown cataphylls and a collar of fibres. Corm 12-20 mm

diam., tunics of medium-textured fibres. Leaves usually

4 or 5, lower 3 or 4 with sword-shaped to falcate blades,

± 1/2 as long as stem and (2-)3-5 mm wide, uppermost

leaf entirely sheathing, main veins and margins slightly

to moderately thickened. Spike flexuose, erect, 2-ranked,

mostly 6-12-flowered; bracts membranous, translueent,

4-5 mm long, outer with prominent central vein or sec-

ondary veins equally prominent, 3-lobed with central

lobe largest or ± equally 3-lobed, inner slightly shorter

than outer, 2-veined and 2-toothed. Flowers nodding

with tepals held vertically, deep pink (sometimes called

purple), white (sometimes edged with dark pink) at base

of tepals, unscented; perianth tube filiform but expanded

near mouth, 3-5 mm long; tepals spreading, oblong,

(8-) 10- 14 X 4—8 mm. Stamens symmetrical in bud,

becoming unilateral in open flower, straight, held hori-

zontally at right angles to tepals; filaments inserted at

mouth of tube, filiform, 2-3(^) mm long, white; anthers

oblong-linear, 2. 5^.0 mm long, yellow, dehiscing com-

pletely by longitudinal slits. Style dividing between base

and lower 1/3 of filaments, style ± 2 mm long, branches

tubular, strongly falcate, stigmatic apically, white. Flow-

ering time: mid-Aug. to late Sept. Figure 5.

Distribution and habitat: Ixia curvata has a relatively

modest range for sect. Dichone, extending in the west-

ern Karoo from the eastern end of the Hantamsberg at

Moordenaarspoort northeast of Calvinia southward to

Middelpos on the Roggeveld Plateau (Figure 1). The

range remains poorly documented and there are no col-

lections from Middelpos itself although we have seen

the species on the commonage there. Our description is

based largely on the Middelpos population and the illus-

tration here is from this population. Plants favour the

red clay soils derived from dolerite but are also found

in pale clay ground derived from Karoo shales. Pollina-

tion of 7. curvata has not been studied. Raimondo et al.

(2009) regarded the conservation status as of Least Con-

cern (LC) and we concur, noting the relatively modest

range of the species.

96

Bothalia 42,2 (2012)

FIGURE 5. — Ixia curvata, Middelpos, without voucher, Scale bar: 10

mm. Artist: John Manning.

Diagnosis and relationships: relatively low grow-

ing, Ixia curvata is a small plant, seldom exceeding 200

mm in height (although taller plants are known) and has

nodding, pink flowers with tepals held vertically with

stamens borne horizontally. Like other members of ser.

Euanthera, the anthers split completely. The four or

five leaves are typically falcate and leathery rather than

fibrotic. Its immediate relationships remain uncertain.

Other species of the series have upright flowers with

tepals spreading horizontally. The plants are notable in

the thick collar of fibres around the base and chestnut-

brown cataphylls.

The species was first collected by Rudolf Marloth in

October 1920, but the label data localizing the plants

from ‘plains near Tulbagh’ is an obvious mistake. Mar-

loth collected in the Roggeveld and nearby in October

1920 when he also made the first collections of Ixia

rigida (initially assigned to 7. brevituba) and I. trifolia,

thereby establishing the presence of sect. Dichone in the

Western Karoo.

Representative specimens

NORTHERN CAPE. 3119 (Calvinia): lower clay slopes of the

Hantamsberg N of Downes Siding, (-BD), 12 Sept. 2004, Goldblatt

& Porter 12411 (MO); 4.5 miles [± 6.8 km] SSE of Calvinia, (-DB),

21 Sept. 1955, Acocks 18504 (PRE). 3120 (Williston): 35 km SE of

Calvinia to Middelpos on Blomfontein road, (-CA), 30 Sept. 1976,

Goldblatt 4281 (MO). Uncertain localities: ‘Plains near Tulbagh,’ Oct.

1920, Marloth 9936 (PRE); Van Rhyns Pass, 15 Sept. 1953, Taylor

3967 (NBG).

6. Ixia trinervata (Baker) G.J.Lewis in Journal

of South African Botany 28: 169. (1962); De Vos: 75

(1999). Tritonia trinervata Baker: 191 (1892); Baker:

212 (1896). Type: South Africa, [Western Cape], Appels-

kraal, Riviersonderend, Ecklon & Zeyher Irid. 242 (K,

holo.!, G, Z, iso.).

Plants 250^50 mm high, stem usually unbranched

or with a single short branch, with membranous cata-

phylls enclosing base. Corm 9-18 mm diam., tunics of

firm, medium-textured, netted fibres. Leaves 2, lower 1

± basal, upper 1 sheathing stem in lower 1/2, blades lan-

ceolate, 4-10 mm wide, conspicuously 3-veined, main

vein moderately thickened, secondary veins slightly

less so, margins not or barely thickened. Spike flexuose,

mostly 7-122-flowered; bracts membranous, translu-

cent, sometimes turning rusty brown in upper 1/3, outer

5-6 mm long, with prominent central vein and up to

six secondary veins, 1 -toothed or ± fringed, inner ± as

long as outer, 2-veined and 2-toothed. Flowers rotate,

deep rose pink, unscented; perianth tube filiform, 4-5

mm long, clasping style; tepals spreading, ovate-ellip-

tic, obtuse to retuse, 12-15 x 6-8 mm. Stamens sym-

metrically arranged, erect; filaments inserted at mouth

of tube, filiform, 2-4 mm long, ± white; anthers erect,

suborbicular, ± 2 mm long, yellow, thecae acute and

recurved at base, dehiscing by narrow, longitudinal

slits almost to apex. Style dividing at base of filaments,

branches falcate, 2. 5-3.0 mm long, tubular-filiform, stig-

matic apically, pale pink.

Distribution and habitat: a narrow endemic of the

Caledon District, Ixia trinervata extends from Elgin Sta-

tion and the Farm Arieskraal in the west to Rivierson-

derend in the east, a linear distance of some 65 km (Fig-

Bothalia42,2 (2012)

97

ure 3). Plants favour clay soils and grow in renosterveld,

often in stony ground. There is evidence that flowering

is stimulated after fire and we have seen plants bloom-

ing well on the Farm Vleitjies after a summer bum. We

agree with Raimondo et al.'s (2009) estimate of near

threatened status (NT) for I. thnervata.

Diagnosis and relationships: first described by J.G.

Baker (1892) as Tritonia trinervata and transferred to

Ixia by G.J. Lewis (1962), /. trinervata is readily rec-

ognized by the presence of just one foliage leaf, the

blade narrowly sword-shaped and with three ± equally

conspicuous veins and unthickened margins. A second

leaf is largely to entirely sheathing, although relatively

broad. The flowers are an intense, bright mauve pink or

red-magenta. Lewis (1962) considered the species most

closely related to usually pale pink-flowered I. stricta,

which blooms later in the season, at which time its 3-5

leaves are usually dry. Both species favour clay or clay

loam soils. The new /. bifolia, described here, recalls I.

trinervata in its single foliage leaf, but the leaf blade is

much narrower than in I. trinervata, with a single main

vein embedded in the leathery-succulent leaf tissue.

When dry the leaf margins appear particularly thickened,

unlike those of /. trinervata. The flowers of /. bifolia are

slightly smaller than those of I. trinervata and have unu-

sual, almost straight, deep pink style branches in marked

contrast to the pale pink, falcate style branches of /. trin-

ervata.

Representative specimens

WESTERN CAPE. — 3419 (Caledon): Diepklowe Nature Reserve,

14 km N of Botrivier, Farm Welgemoed, (-AA), 3 Sept. 2001, Helme

2152 (NBG); Elgin, Die Hawe, near Post Office, clay ground, (-AA),

11 Oct. 2000, Goldblatt 11639 (MO); Lebanon Forest Reserve, Houw

Hoek, (-AA), 26 Sept. 1954, Martin 1043 (NBG), Lewis 4178 (PRE,

SAM); slopes of Eseljag Pass, near Farm Vleitjies, clay soil in renos-

terveld, (-AB), 30 Sept. 2000, Goldblatt & Ndnni 11597 (MO, PRE);

18 Sept. 2011, Goldblatt & Manning 13642 (MO, NBG).

7. Ixia bifolia Goldblatt & J.C. Manning, sp. nov.

TYPE. — Western Cape, 3419 (Caledon); northern

slopes below Shaw’s Pass, Farm Treintjiesrivier, (-AD),

13 Sept. 2009, Goldblatt & Manning 13444 (NBG,

holo.; MO, PRE, iso.).

Plants mostly 160-300 mm high, stem unbranched,

with short scale ± 1 mm long, inserted 12-30 mm below

base of spike, sheathed basally by dry, brown cataphylls.

Corm 10-12 mm diam., tunics of fine, soft to firm fibres

extending upward as collar, bearing cormlets at base.

Leaves 2, lower Iwith long sheath, ± lanceolate, blade

60-80 X (3-)5-8 mm, reaching up to middle of stem,

leathery-succulent, main vein not raised when alive (vis-

ible as a pale line when held to light), slightly raised

when dry, margins thickened, hyaline, transparent when

alive, when dry raised above mesophyll and yellow,

upper leaf largely sheathing, sometimes with short free

blade similar to basal but 2-3 mm wide. Spike mostly

4— 7-flowered, ± straight; bracts translucent pink with

purple veins, 5-6 mm long, outer with 3 main veins ter-

minating in acute teeth and 1 or 2 secondary veins along

margins, inner 2-veined and acutely 2-toothed. Flowers

rotate, upright, deep pink, darker toward tepal bases,

rose-scented; perianth tube filiform, 5-6 mm long;

tepals obovate, 12-14 x 5-7 rnm. Stamens symmetri-

cally arranged; filaments filiform, inserted at mouth of

narrow part of tube, ± 1 .5 mm long, deep pink (drying

purple); anthers suborbicular, 2.0-2. 5 mm long, yellow,

thecae obtuse basally, dehiscing completely or almost so

by narrow slits. Style dividing at base of filaments, style

branches 3. 8^.0 mm long, deep pink (drying purple),

± straight, extending outward below anthers, expanded

and bilobed at tips. Capsules globose, ± 4 mm diam.

Flowering time: mostly late Aug. to mid-Nov. but Dec.

and early Jan. in the Caledon Swartberg. Figure 6.

Distribution and habitat: restricted to the Caledon

District, Ixia bifolia is known from the northern slopes

of Shaw’s Mtn south of Caledon and from the lower

slopes of the Swartberg east of Caledon (Figure 3). The

soil at the Shaw’s Mtn site is hard, stony clay and the

habitat there is shared by the rare Tritoniopsis flexuosa

(L.f ) G.J.Lewis (known from just two other localities).

With its narrow range and just two known populations,

one small and near a major road, we suggest a conserva-

tion status of (VU), Vulnerable.

Diagnosis and relationships: Ixia bifolia seems at

first to belong in the I. micrandra-confusa complex,

which also has very slender stems and small flowers but

it stands out in the very short, leathery-succulent foli-

age leaf blade, up to 80 mm long and 5-7 mm wide.

The leaf has a single, prominent vein and moderately

thickened margins. The flowers seem most like those of

I. confusa but the short, deep pink filaments ±1.5 mm

long and similarly deep pink, straight style branches ±

4 mm long (both drying purple) are quite different from

I. confusa, which has white filaments 1.3-2. 5 mm long

and white, strongly falcate style branches 1-2 mm long.

We suspect that /. bifolia may be more closely allied to

/. trinervata, which has slightly larger flowers on longer

spikes but also a single foliage leaf, this relatively broad

and always with three prominent veins and margins not

thickened. I. bifolia shares with other members of the /.

micrandra complex the finely fibrous corm tunics that

extend upward around the stem base. Stems are almost

always unbranched and have an odd feature, a small,

nearly microscopic, translucent scale and prophyll, ±

1 mm long, inserted 12-30 mm below the base of the

spike. Stems of I. confusa may be branched and are nor-

mally taller than in I. bifolia but also have a translucent

stem scale in populations from the Swartberg and some-

times elsewhere across its range. /. confusa has 3 or 4

linear leaves, usually ± 2 mm, but occasionally to 4 mm

wide.

Representative specimens

WESTERN CAPE. — 3419 (Caledon); N slopes below Shaw’s Pass

E of main road, (-AD), 16 Sept. 2009, Goldblatt & Manning 13640

(MO, NBG, K); lower slopes of Caledon Swartberg, grazed field E of

Drayton Siding, (-BA), 16 Sept. 2011, Goldblatt & Manning 13641

(MO, NBG).

8. Ixia confusa (G.J.Lewis) Goldblatt &

J. C. Manning, stat. nov. I. micrandra var. confusa

G.J.Lewis: 162 (1962); De Vos: 73 (1999). Type: South

Africa, [Western Cape], Montagu, Donkerkloof, 26 Sept.

1946, Compton 18481 (NBG, holo.!; BOLl, iso.).

Plants (150-)250-500 mm high, stem simple or

1 -branched, with short scale ± 1 mm long, inserted

98

Bothalia 42,2 (2012)

FIGURE 6, — Ixia hifolia, Goldhialt & Manning 13444 (NBG). A, flow-

ering plant; B, stamens and style branehes. Scale bar: 10 mm.

Artist: John Manning.

25^0 mm below base of spike, sheathed basally by

green, leathery to firm, dry cataphylls. Corm 8-10 mm

diam., tunics of fine, soft fibres sometimes extending

upward in a weakly developed collar. Leaves (2)3(4),

lower (1)2 or 3 with linear to narrowly sword-shaped

blades, ± 1/3 as long to reaching top of stem, l.S^.O

mm wide, upper leaf largely sheathing, with short free

blade. Spike slightly flexuose, (2-)5-9(-12)-flowered;

bracts membranous, becoming dry from tips, 4-5 mm

long, outer 1- or 3-veined, irregularly 2-3(-5)-lobed or

shortly toothed, inner often slightly longer than outer,

2-veined and 2-toothed. Flowers rotate, upright, pink

to light red-purple, rose-scented; perianth tube filiform,

3^ mm long; tepals spreading, ± ovate, mostly 12-15

X 6-7 mm. Stamens symmetrically arranged, erect; fila-

ments inserted at mouth of tube, filiform, 1.0-2. 8 mm

long; anthers oblong, 2. 0-2. 3 mm long, yellow, thecae

acute and recurved at base, usually dehiscing longitu-

dinally from base almost to apex. Style dividing at or

up to middle of filaments, style branches 1. 5-2.0 mm

long, falcate, tubular with expanded ciliate, stigmatic

tips, sometimes shortly forked apically. Flowering time:

mostly Jul. -early Dec.

Distribution and habitat'. Ixia confusa occurs in the

mountains of southern Western Cape, from near Elim

and the Riviersonderend Mtns in the west through the

Langeberg to the Paardevlei Mtns near Farm Bonnie-

vale at the western end of the Outeniqua Mtns {Vlok 709

NBG, PRE), and inland in the Great Swartberg (Figure

7) where it has been recorded from Swartberg Pass,

Meiringspoort, and Blesberg slightly further east. An

outlying collection {Marloth 2429 PRE) from Montagu

Pass near George extends the range slightly south. We

have encountered plants {Goldblatt & Manning 12946)

on moist southwest-facing slopes in loamy clay in renos-

terveld-thicket transition vegetation. The flowers of our

collection were strongly rose-scented. We suspect that

plants flower particularly well after fire but our col-

lection was made in veld that had not burned for over

10 years. Occurring mostly in montane habitats, there

seems no current threat to the species and we agree with

Raimondo et al. ’s (2009) (LT, as I. micrandra var. con-

fusa), Least Threat conservation status.

Diagnosis and relationships', known to G.J. Lewis

(1962) as Ixia micrandra var. confusa, we raise the

taxon to species rank after examining living plants and

16° 18^ 20° W° 24° 26° ^

FIGURE 7. — Distribution of Ixia confusa, ▲; I. micrandra, o; and I.

minor, •.

Bothalia 42,2 (2012)

99

comparing them with I. micrandra. Both I. micrandra

and closely related I. minor (= /. micrandra var. minor

G.J. Lewis) have corms with finely fibrous tunics that

extend upward in a collar around the base of the stem,

terete to subterete or linear leaves mostly 1.0- 1.5 mm

at widest, and spikes of up to 6 flowers. /. confusa has

corms with similar finely fibrous tunics, sometimes

without a collar of fibres around the base, but plane

leaves l.S^.O mm wide, and spikes with as many as

12 flowers. I. micrandra has anthers 3-4 mm long,

and slender, ± straight, ascending style branches 3^

mm long, thus quite different from the shorter anthers,

± 1.0-2. 5 mm long, and falcate style branches 1.5-2. 5

mm long of I. confusa. The style branches of /. confusa

match those of I. minor but the terete or subterete leaf

blades, flattened filaments and consistently few-flowered

spikes of the latter immediately separate them.

I.xia confusa may also be confused with the new I.

bifolia but this species has a single, relatively broad,

leathery-succulent foliage leaf and flowers with shorter

filaments ± 1 mm and longer, straight style branches ±

4 mm long. Both the filaments and style branches of /.

bifolia are dark pink and dry dark purple, unlike the pale

filaments and style branches of I. confusa, which dry to

a pale, almost colourless state.

Annotations by G.J. Lewis on collections of Ixia con-

fusa from the Swartberg indicate that at one time she

considered these populations a separate species, which

she intended to call 1. aestivalis, in part because of their

late flowering, from late October to early December.

We find that collections from the Swartberg also differ

in having slightly shorter filaments, 1-2 mm long, than

typical I. confusa (1.5-2. 8 mm long), the style some-

times dividing above the mouth of the perianth tube

(opposite the middle of the filaments in Goldblatt 7962),

and anther dehiscence sometimes not reaching the mid-

dle of the thecae. The plants have a small, translucent

scale and prophyll on the upper part of the stem, a fea-

ture occasionally present in southern populations of I.

confusa. Additional study may show that the later-flow-

ering Swartberg populations merit separate taxonomic

status but we find no consistent character differences

between them.

Representative specimens

[* = apparently no collar around base of stem]

WESTERN CAPE. — 3319 (Worcester): eastern foot of Stettyns-

berg, (-CD), 31 July 1949, Eslerhuysen 15590 (BOL, NBG, PRE);

slopes of Boesmanskloof Pass, MacGregor, (-DC), 13 Sept. 1962,

Lewis 6067* (NBG, PRE); top of Boesmanskloof Pass, MacGregor,

(-DC), 13 Sept. 1962, Lewis 6080 (NBG). 3320 (Montagu): foot of

Tradouw Pass, (-DC), 18 Sept. 1968, Marsh 875 (PRE). 3321 (Ladi-

smith): Paardevlei Mtns N of Farm Bonniedale, 1 500 ft [± 460 m],

(-DD), 24 Sept. 1983, Vlok 709* (NBG, PRE). 3322 (Oudtshoom):

summit, Swartberg Pass, (-AC), Dec. 1906, H. Bolus 12336 (BOL);

Swartberg Pass, (— AC), 5 Nov. 1958, Werdertnann & Oberdieck 843

(B, PRE), 22 Oct. 1986, Goldblatt 7962 (MO, PRE); Swartberg, sandy

flats SW of Blesberg, (-BC), 7 Jan. 1975, Thompson 2274 (PRE);

Meiringspoort, S slopes, (-BC), 16 Oct. 1955, Esterhuysen 24880

(BOL, MO, PRE); Montagu Pass, George, (-CD), Marloth 2429

(PRE). 3419 (Caledon): Uintjieskuil near Elim, (-DB), Sept. 1926,

Smith 3182* (PRE). 3420 (Bredasdorp): S of Swellendam on road

to Bontebok Park, light stony clay, (-AB), 26 Aug. 2000, Goldblatt

11430 (MO); coastal renosterveld hills at Goerreesoe, S slope, (-AD),

30 Aug. 1962, Acocks 22672 (PRE); clay loam slopes between Heidel-

berg and Strawberry Hill, in renosterveld, (-BB), 4 Sept. 2007, Gold-

blatt & Manning 12946 (MO, NBG, PRE), 3421 (Riversdale): near

Oudebosch crossing, Korente River crossing, Riversdale, (-AA), 16

Sept. \99,\,Hugo 2 773 (PRE).

9. Ixia micrandra Baker in Journal of Botany 1876;

237 (1876); Baker: 78 (1896); Lewis: 159 (1962); De

Vos: 71 (1999). Type: South Africa, [Western Cape],

Houw Hoek Mtns, July 1830, Zeyher 4011 (K, lecto.!

designated by Lewis: 161 (1962); PRE!, SAMI, S, Z,

isolecto.).

Plants 250-350 mm high, stem unbranched, base

sheathed by membranous to ± dry, brown cataphylls.

Corm 9-18 mm diam., tunics of fine fibres extending

upward as a collar. Leaves 2, lower 1 ± basal, upper 1

sheathing the stem in lower 1/2 to 2/3, free distally for

up to 100 mm, blades linear, (0.5-)1.0-1.5(-2.0) mm

wide, margins and midrib lightly thickened, hyaline

when dry. Spike erect, ± straight, (l-)2^(-6)-flowered;

bracts membranous, translucent, sometimes suffused

with purple distally, outer 5-7 mm long, prominently

3-veined and usually with secondary veins evident,

acutely 3 -cuspidate, sometimes central cusp smaller,

inner ± as long as outer or slightly longer, 2-veined

and 2-cuspidate. Flowers rotate, pale mauve-pink; peri-

anth tube filiform, 3-4 mm long, clasping style; tepals

spreading, ovate-elliptic, 12-15 x 4—6 mm. Stamens

symmetrically arranged, erect; filaments inserted at

mouth of tube, linear, straight, 0.6-1.8(-2.5) mm long,

± pale pink; anthers oblong, 2. 0-2. 6 mm long, erect, yel-

low, thecae slightly recurved basally and acute dehisc-

ing completely from base by longitudinal slits. Style

dividing at or shortly above base of filaments, branches

± straight and diverging to slightly arched, 2-5 mm

long, usually longer than stamens, filiform-involute,

stigmatic and sometimes forked at apex, pale pink.

Flowering time: mostly late Jul.-early Sept., occasion-

ally later.

Distribution and habitat: extending from Houw

Hoek, Bot River and the Caledon Swartberg to

Bredasdorp and De Hoop (Figure 7), Ixia micrandra

typieally occurs on sandstone slopes and flowers well

only in seasons immediately after veld fires. A col-

lection from De Hoop is from pockets of sandy soil in

limestone pavement, an unusual habitat for the species.

Largely montane, we see no current threat to the species

and endorse Raimondo et al. ’s (2009) (LT), least threat,

status.

Diagnosis and relationships: treated by G.J. Lewis

(1962) as including the two additional varieties, var.

confusa and var. minor, we have no doubt that /. micran-

dra is allied to these two taxa, which are here raised to

species rank. I. micrandra stands out in the group in

its long, narrow, almost linear filaments and its style

branches, which are unusually long at 2. 0-3. 5 mm, and

straight or slightly arched, thus extending between the

bases of the anthers. /. confusa has similarly slender but

somewhat shorter filaments, slightly shorter anthers,

and shorter, strongly arched style branches, 1. 5-2.0

mm long. The leaves of I. confusa have plane, usually

wider blades up to 4 mm wide (vs. 0.5-1.5(-2.0 mm)

and without the prominently thickened main vein of M.

micrandra. The style branches are also strongly arched

in I. minor but that species has unusual flattened, Ian-

100

Bothalia 42,2 (2012)

ceolate filaments, narrowed and kinked at the tips. The

ranges of I. micrandra and I. minor are almost eomple-

mentary, that of I. micrandra extending east of I. minor,

but the two have been collected growing and flowering

on the same day in the Houw Hoek Mtns, where their

distributions overlap.

The only significant variant of Ixia micrandra we

have seen is from near Bot River [MacOwan in Herb.

Norm. Austr. Afr. 262). Plants of this population have

unusual thick, fleshy leaf blades, but the flowers and

other features accord with the species.

Representative specimens

WESTERN CAPE. 3419 (Caledon): Houw Hoek, (-AA), 17 Sept.

1987, [collected with I. minor, Beyers 17, on same day], Beyers 22

(NBG); Bot River, Langhoogte, (-AA), Aug., Ecklon & Zeyher Irid.

92 (MO); Caledon Baths, (-AB), July 1892, F. Guthrie 2531 (BOL);

Caledon Swartberg and vicinity of the baths, (-AB, BA), August

1830, Zeyher 4009 (PRE); Highlands, low slopes between Kleinmond

and Bot River (-AC), 27 July 1960, Lewis 5733 (NBG); raised plain

between the mountains and mouth of the Bot River, (-AC), July 1885,

MacOwan in Herb. Norm. Austr. Afr. 262 (BOL, PRE); Kleinmond

commonage, (-AC), 15 Aug. 1982, Burman 852 (BOL); Hermanus,

behind Golf Course, deep sand, 60 m, (-AD), 1 Aug. 1984, 5. Wil-

liams 1042 (MO); Caledon Swartberg, well drained sandy soil, (-BA),

1 Aug. 2004, Raimondo CR278 (NBG); Genadendal, mountains at ±

4 500 ft [1 400 m], (-BA), 27 Oct. 1899, Galpin 4686 (BOL), SaL

monsdam Reserve, Stanford, burnt slopes, 1 000 ft, (-BB), 31 Aug.

Goldblatt 421 (BOL); 2 miles [3 km] NW of Papiesvlei P.O., (-BC),

25 July 1962, Acocks 22449 (PRE); Bredasdorp, hillside, (-BD), 27

July 1955, Van Niekerk 430 (BOL). 3420 (Bredasdorp): De Hoop, Pot-

berg Nature Reserve, upper slope above Boskloof, back sandy ground,

(-BC), 6 Sept. 1978, Burgers 7067 (PRE).

10. Ixia minor (G.J.Lewis) Goldblatt & J.C. Manning,

stat. nov. /. micrandra var. minor G.J.Lewis: 162 (1962);

De Vos: 73 (1999). Type: South Africa, [Western Cape],

Hottentots Holland Mtns, 10 Sept. 1928, J. Hutchinson

481 (BOL, holo.!; K, PRE!, iso.).

Plants 200-300 mm high, stem unbranched, sheathed

by membranous to dry, brown cataphylls at base. Corm

8-10 mm diam., tunics of fine fibres extending upward

as a collar, often in a dense mass. Leaves 2, lower 1 ±

basal, upper 1 sheathing stem in lower 1/2 to 2/3, free

distally for short distance, blades ± terete or oval in sec-

tion, up to 1 mm diam., main vein evident only when

dry. Spike erect, straight, mostly 2-4-flowered, bracts

membranous, translucent, usually flushed purple, outer

5-7 mm long, usually prominently 3-veined, shallowly

3-lobed to acutely 3-toothed, inner bract ± as long as

or slightly longer than outer, 2-veined and 2-lobed or

2-toothed. Flowers rotate, bright pink to mauve pinkj

perianth tube filiform, 3-5 mm long, clasping style;

tepals spreading, ovate-elliptic, 12-14 x 5-6 mm. Sta-

mens symmetrically arranged, erect, filaments flattened,

± lanceolate but narrowed and kinked at tips, inserted

at mouth of tube, ± white, 1.0(-L5) mm long, anthers

ovoid, ± 1.0 mm long, erect, yellow, thecae acute but

abruptly incurved and touching at base, dehiscing fully

by longitudinal slits, often more widely at base, yel-

low. Style dividing opposite base to middle of filaments,

branches up to 2 mm long, involute, tubular-falcate, ±

compressed, narrowed at tips, stigmatic apically, white.

Flowering time: late Jul.-mid-Sept., rarely later.

Distribution and habitat', a narrow endemic of the

southwestern Cape, Ixia minor occurs from Franschhoek

Pass and the Wemmershoek Mtns south though the Hot-

tentots Holland and Kogelberg Mtns to Houw Hoek

(Figure 7). The habitat, stony sandstone slopes, is the

same as that for I. micrandra and as in that species we

see no current threat and suggest a least threat (LT) con-

servation status.

Diagnosis and relationships'. Ixia minor, treated as

var. minor of I. micrandra by Lewis (1962) but raised

here to species rank, is evidently closely allied to the

more widespread I. micrandra, with whieh it is easily

confused unless floral details are examined carefully.

Both share a slender, usually unbranched stem, small

corms with fine, netted tunics that extend upward in

a collar around the base, and a dry, often brown, upper

cataphyll. The critical differences between the two spe-

cies lie in their flowers: I. minor has flattened, ± wedge-

shaped to oblanceolate filaments abruptly narrowed at

the tips, shorter and broader anthers, often not dehiscing

to the apex and kinked at the base with the bases of the

thecae turned inwards and touching, and shorter, wider,

strongly arched style branches slightly compressed and

narrowed at the tips. To these microscopic details can

be added a deep pink perianth. Flowers of I. micran-

dra have pale pink to purple-pink tepals, linear fila-

ments, longer, oblong anthers dehiscing to the apex and

longer, more slender style branches, ± straight to slightly

arched.

The leaves of the two species also differ: those of

Ixia minor are subterete to oval in section, thus when

alive without margins or evident central vein, whereas I.

micrandra h^ plane, linear leaf blades with somewhat

thickened margins and, when dry, a raised main vein.

Floral differences between I. minor and I. micrandra are

illustrated in detail in the accounts by both Lewis (1962)

and De Vos (1999). A color photograph of I. minor

(Manning et al. 2002: 242) under the name I. micrandra

clearly shows the short, recurved style branches of the

species. Compelling evidence for their separation is pro-

vided by collections of both species made by the late J.

Beyers in the Houw Hoek Mtns flowering on the same

date and sufficiently different, in appearance that Beyers

gave them different collection numbers. Habitat differ-

ences, if any, were not recorded.

Representative specimens

WESTERN CAPE. — 3319 (Worcester): Wemmershoek, SE end,

3 000-4 000 ft [915-1 220 m], (-CC), 19 Oct. 1943, Esterhuysen 9060

(BOL); Franschoek Pass, ± 1 km S of summit, (-CC), 9 Oct. 1973,

Boucher 2337 (PRE). 3418 (Simonstown): W foot of Sir Lowry’s Pass,

(-BB), Aug. 1985, De Vos 2585 (NBG); Sir Lowry’s Pass mountains,

(-BB), Sept. 1894, H. Bolus 7850 (BOL); Steenbras River, 600 m, (-

BB), 14 Oct 1894, Schlechter 5402 (PRE). 3419 (Caledon): Elgin, (-

AA), 19 Sept. \93\,Levyns 3341 (BOL); Hottentots Holland, Moorde-

naarskop and Sugar Loaf, (-AA), 30 Oct. 1943, Esterhuysen 9123

(BOL); Houw Hoek, (-AA), 17 Sept. 1987 [collected with 7. micran-

dra, Beyers 22, on same day], Beyers 17 (NBG).

Ser. Dichone

Flowers upright or nodding, radially symmetric

{Ixia stricta) or zygomorphic. Stamens symmetrically

arranged {Ixia stricta) or unilateral and horizontal to

pendent; anthers oblong, recurved and acute at base,

dehiscing incompletely from base. Style branches always

falcate.

Bothalia 42,2 (2012)

101

7 spp., centred in the western half of Western Cape,

but with isolated populations in northern Namaqualand

and the Bokkeveld Mtns of Northern Cape.

11. Ixia stricta (Eckl. ex Klatt) G.J. Lewis in Jour-

nal of South African Botany 28: 167 (1962); De Vos:

73 (1999). Tritonixia stricta Eckl. ex Klatt: 357 (1882).

Tritonia scillaris var. stricta (Eckl. ex Klatt) Baker:

191 (1892). Type: Western Cape, Caledon, Swartberg

and vicinity of the Baths, December 1830, Zeyher 4008

(SAM, lecto.!, designated by Lewis: 162 (1962); K,

PRE!, isolecto.).

Agretta stricta Eckl.: 23 (1827), nom. nud.

Plants 200^50 mm high, stem simple or with 1 or 2

± erect branches subtended by acute, translucent bracts

and prophylls, sheathed at base by short, dry cataphylls.

Corm mostly ± 10-15 mm diam., tunics of medium-

textured, shaggy fibres. Leaves (3)4 or 5, often ± dry

at flowering, basal, lower 2 or 3 with blades plane, 4-9

mm wide, upper 1 or 2 sheathing stem in lower 1/2 to

2/3, free distally for short distance, margins and main

vein thickened (hyaline when dry). Spike erect, flexu-

ose, 6-12(-18)-flowered, bracts membranous, translu-

cent, outer 5-6 mm long, usually with 1 prominent vein,

and 1 -toothed, inner bract ± as long or slightly shorter

or longer, 2-veined and 2-toothed. Flowers rotate, pink

to purple, pale yellow edged with darker pink at tepal

bases, sweetly scented (Viviers 1150), perianth tube

filiform, 4—5 mm long, clasping style, tepals spread-

ing, obovate, ± obtuse, ± 10 x 4-5 mm. Stamens sym-

metrically arranged, filaments 1.0- 1.5 mm long, white,

anthers oblong, ± 2 mm long, erect, yellow, thecae acute

and slightly recurved at base, dehiscing near base by

longitudinal slits. Style dividing opposite base of fila-

ments, branches arching outward, ± 1.5 mm long, fal-

cate, tubular, stigmatic apically, pale pink. Flowering

time: Nov. -mid-Dec.

Distribution and habitat: restricted to the Caledon-

Bredasdorp area of Western Cape, Ixia stricta extends

from the Houw Hoek Mtns in the west to Bredasdorp in

the southeast (Figure 1). Typically flowering late in the

year, in November and December, I. stricta is poorly

collected. Not surprisingly the foliage is often more-or-

less dry at flowering. Plants occur mostly in montane

habitats, but usually at fairly low elevations, in sand-

stone and loamy clay soils. Raimondo et al. (2009) sug-

gest a conservation status of (LC), least concern, but we

suggest that (NT), near threatened, is more appropriate.

Diagnosis and relationships: while not particularly

distinctive in its radially symmetric flowers with short,

erect stamens, Ixia stricta is identified in ser. Dichone

by a combination of the rotate flowers and a basal fan of

(3)4 or 5 sword-shaped leaves, the blades with a promi-

nent main vein and thickened margins. The sweetly

scented flowers have a filiform tube about half as long

as the tepals and a particularly short style ± 1 mm long.

Blooming late in the season, in November and Decem-

ber, the leaves are often dry or partly so, a feature unu-

sual in the genus.

History: the plant illustrated by Jacquin (1792)

and misidentified as Ixia polystachya L. is the earli-

est record of Ixia stricta and this illustration is also one

of two sources cited by Ker Gawler (in Sims 1801) for

his I. scillaris var. angustifolia. Both Lewis (1962) and

De Vos (1999) associated Jacquin’s illustration with I.

stricta and the illustration shows clearly the relatively

broad, prominently veined basal leaves and the sheath-

ing upper leaves subtending straight, several-flowered

branches. Ker Gawler’s varietal name angustifolia is

nomenclaturally illegitimate and superfluous because he

included the earlier name I. scillaris var. incarnata, this

now most likely I. scillaris subsp. lat folia.

Watsonia retusa (Salisb.) Klatt (1882), a combina-

tion based on indirect citation of the basionym Ixia

retusa Salisb., was regarded by Lewis as a synonym of I.

micrandra var. confusa. De Vos (1999), however, asso-

ciated the species with /. stricta and we concur. Klatt

evidently intended to provide a name for the plant called

I. polystachia [sic] by Ker Gawler (1803) in Curtis’s

botanical magazine but as Ker Gawler specifically men-

tioned I. retusa Salisbury (1796) in his discussion of the

plate in question, Klatt’s name must be treated as a com-

bination, but with a basionym that has no known type.

Salisbury’s brief protologue matches the plant depicted

in Curtis’s botanical magazine published seven years

later in all respects, notably the style branches emerging

between the filaments and the tepals twice as long as the

perianth tube. But there is no connection between Salis-

bury’s species and the painting.

Representative specimens

WESTERN CAPE.— 3419 (Caledon): Houw Hoek, 2 000 ft [610

m], (-AA), 25 Nov. 1896, Schlechter 9411 (BM, BOL, K, PRE);

Groenland Mtns, Klein Jakkalsrivier, red clay, 2 200 ft [± 700 m], (-

AA), 2 Dec. 1982, Viviers 1150 (NBG, PRE); Hermanus, Glenfivin,

Onrus Mtn, fynbos on clay, recently burnt, (-AC), 16 Nov. 1991,

Barker 305 (MO); Vogelgat, Hermanus, at Femkloof gate, on shale

band, 450 m, (-AD), 27 Nov. 1983, Williams 3523 (NBG, PRE);

Drayton Siding, Caledon, (-BA), 16 Dec. 1968, Goldblatt 393 (BOL);

roadside between Napier and Elim, (-BD), 5 Dec. 1985, De Vos 2642

(NBG); near Napier, (-BD), 30 Nov. 1933, Salter 4151 (BOL, K);

near Baardscheerdersbos, (-DA), 10 Nov. 1985, De Vos 2636 (NBG);

Boskloof, Bredasdorp, (-DB), 28 Dec. 1983, Lavranos 21797 (MO,

PRE); Bredasdorp, Farm Geelrug, (-DB), 17 Dec. 1975, Barker 10940

(NBG, PRE). 3420 (Bredasdorp): Bredasdorp Mtns, (-CA), Galpin

s.n. PRE10463 (PRE).

12. Ixia erubescens Goldblatt in Journal of South

African Botany 37: 233 (1971b), as a new name for 1.

crispa L.f: 91 (1782), a superfluous name for I. undu-

lata Burm.f Agretta crispa Eckl.: 24 (1827), as a new

name for /. crispa L.f. Tritonia thunbergii N.E.Br.: 137

(1929), as a new name for Ixia crispa in Tritonia, non T.

crispa (L.f.) Ker Gawl. Type: South Africa without pre-

cise locality, Thunberg s.n. [LINN 58.25, lecto., desig-

nated by Lewis: 173 (1962)].

Plants 120-300 mm high, usually forming clumps,

stem simple, rarely 1 -branched, sheathed basally by

membranous cataphylls. Corm mostly 5-8 mm diam.,

tunics of fine fibres, usually with large cormlets at base.

Leaves 4-6, ± lanceolate, all basal, ± 1/3 to 1/4 as long

as stem, 2-4(-6) mm wide, main vein well developed,

margins or half blade crisped and deeply undulate. Spike

erect, lax, secund, weakly flexuose, mostly 6-10-flow-

ered, flowers mostly 15-25 mm apart; bracts mostly

green, becoming translucent at tips, turning purple

along distal margins, 3-^ mm long, outer with 3 acute-

102

Bothalia 42,2 (2012)

attenuate cusps, obscurely 3-veined, inner 2-veined and

acutely 2-toothed. Flowers ± second with tepals held ±

vertically, zygomorphic with stamens unilateral, pink,

yellow-green in throat edged with dark purple at tepal

bases, unscented; perianth tube filiform, 2-3 mm long,

tightly clasping style; tepals ovate, spreading ± at right

angles to tube, 8-11 x 4-6 mm, outer tepals slightly nar-

rower than inner. Stamens unilateral; filaments filiform,

± 2 mm long, greenish white, drying yellow; anthers

oblong, ± 1.5 mm long, yellow, horizontal, becom-

ing slightly pendent, thecae acute and recurved at base,

dehiscing incompletely by a narrow slit at base, yellow.

Stymie dividing at base of filaments, style branches 1.5-

2.0 mm long, greenish white, falcate, recurved in distal

third, tubular, stigmatic apically. Flowering time\ mainly

mid-Aug.-mid-Sept.

Distribution and habitat: predominantly a species

of near southwestern Western Cape, Ixia erubescens

extends from Tulbagh southwards to Stellenbosch and

the Caledon District (Figure 8) where its southeastern-

most station is at Franskraal, but is absent from the

Cape Peninsula. Outlying populations occur on the

lower slopes of the Piketberg. The species is most often

encountered on heavy clay or granitic alluvium in sea-

sonally damp places, where plants typically form tight

clusters through the germination of daughter cormlets.

The Piketberg plants are restricted to cooler south-fac-

ing slopes where they grow on well-drained clay slopes

in the shade of shrubs and small trees, and grow singly

rather than in clusters. We concur with the conservation

status of /. erubescens, listed by Raimondo et o/.(2009)

as (LC), least concern, although we note that its range

has been historically much reduced through agriculture,

especially in the Swartland and Boland.

Diagnosis and relationships: a relatively small spe-

cies, Ixia erubescens is immediately recognized by the

fine corm tunics, the 4—6 narrow, deeply undulate and

crisped leaves, usually unbranched stem, and the verti-

cally oriented flower with the stamens unilateral and

slightly pendent when fully open. Both Lewis (1962)

and De Vos (1999) regarded the flowers as fundamen-

tally actinomorphic with symmetrically arranged sta-

mens and noted that only late in anthesis do the anthers

become unilateral and pendulous, simply due to grav-

ity. Our own observations counter this: the stamens

are unilateral immediately the flower buds open. We

FIGURE 8. — Distribution of Ixia colUna, A; I. erubescens, •; and I.

flagel laris, o.

assume that I. erubescens is most closely allied to /. scil-

laris, which typically has medium to coarse corm tunic

fibres, broader leaves, usually with plane margins, and

somewhat to substantially larger flowers, also with uni-

lateral stamens, but with a perianth tube up to 4.5 mm

long and tepals up to 14 mm long (vs. tube 2-3 mm long

and tepals 11-12 mm long). Occasional populations of

I. scillaris with undulate or crisped leaves are readily

distinguished from I. erubescens by their broader leaves,

larger flowers, and coarser corm tunic fibres.

The taxonomic history of Ixia erubescens is some-

what confused. Linnaeus fil. (1782) described I. crispa

based on C.P. Thunberg’s collection but cited the earlier

name I. undulata Burm.f. (Burman 1768) (now Trito-

nia undulata) in synonymy. According to current rules

of nomenclature this rendered I. crispa superfluous and

therefore illegitimate. The two were often confused by

later authors and Baker (1877) used the name Tritonia

undulata for the species, not realizing that its type is

a different plant. Brown (1929) was the first to realize

that /. crispa and I. undulata were different species and

transferred I. crispa L.f to Tritonia under a new epi-

thet, T. thunbergii, because he believed that the name T.

crispa (L.f) Ker Gawl. (based on Gladiolus crispus L.f)

barred a combination based on Linnaeus fil’s I. crispa.

On transferring sect. Dichone to Ixia, Lewis (1962)

used the name I. crispa for the species. Then Goldblatt

(1971b), realizing that 7. crispa was superfluous and thus

illegitimate, established the new name I. erubescens.

Transfer to Ixia of the legitimate T. thunbergii N.E.Br.

is prevented by use of that epithet in Ixia by Roemer

& Schultes (1817), their species being a synonym of T.

securigera Ker Gawl.

Representative specimens

WESTERN CAPE. — 3218 (Clanwilliam): Piketberg, SE-facing

slopes of Versveld Pass, damp clay and clay-loam slopes, (-AC), 6

Sept. 2009, Goldblatl & Porter 13280 (MO, NBG, PRE), 2 Nov. 2011

(fr.), Goldblatt & Porter 1 3712 (MO, NBG). 3318 (Cape Town): near

Wellington, (-DB). 17 Aug. 1926, Grant 2383 (BOL, PRE); stony hills

near Paarl, (-DB), Ecklon 10 'Agretta crispa' (BM, G, M, K, PRE);

Elsenberg, (-DD), Grant 2516 (MO, PRE). 3319 (Worcester): Tul-

bagh, (-AC), Oct. 1916, Marloth 7772 (PRE), Sept. 1915, Marloth

7127 (PRE), 11 Sept. 1926, Grant 2467 (PRE); Tulbaghskloof, etc..

Sept. 1830, Ecklon & Zeyher s.n. 'Agretta crispa' (PRE); near Artois,

Tulbagh, (-AC), Aug. 1885, H. Bolus s.n. (BOL26898); Tulbagh, stony

clay hillside, 12 Sept. 1950, Esterhuysen 17474 (BOL). 3419 (Cal-

edon): Houw Hoek, (-AA), Aug. 1912, Rogers s.n. (PRE 13553); 3 km

W of Caledon, W of road to Badshoogte, (-AB), 12 Oct. 2001, Helme

2294 (NBG); Farm Karwyderskraal, Bot River-Hermanus, (-AC), 15

Sept. 1988, O'Callahan 1784 (NBG); Franskraal, N slope in damp

clay, (-CB), 29 Sept. 2005, Ebrahim & Raimondo CRI075 (NBG).

13. Ixia scillaris L., Species plantarum ed. 2, 1: 52

(1762); Lewis: 173 (1962); De Vos: 78 (1999). Trito-

nia scillaris (L.) Baker: 163 (1877). Tritonixia scillaris

(L.) Klatt: 357 (1882). Type: South Africa, without pre-

cise locality or collector [LINN 58.25 — digital image!,

lecto., designated by Lewis: 174 (1962)].

Plants mostly 150-500 mm high, stem often with

1-2, rarely up to 6 branches, sheathed basally by ±

dry, papery cataphylls, often green distally, later turn-

ing brown. Corm 10-15 mm diam., tunics of medium-

textured to coarse fibres; cormlets at base conspicu-

ous in subsp. toximontana. Leaves 3-7, mostly in basal

fan, upper 1 or 2 partly sheathing the stem, 1/3 to 1/2

Bothalia 42,2 (2012)

103

as long as stem, blade lanceolate or falcate, (3-)6-20

mm wide, main and sometimes additional veins raised

but not thickened, margins markedly thickened in

subsp. toximontana. Spike erect, fairly lax, flexuose to

± straight, secund in 2 ranks, mostly 1 0-20-flowered;

bracts membranous, translucent, sometimes turning pur-

ple distally, 3-5 mm long, outer prominently 3-veined

and 3-toothed, inner 2-veined and 2-toothed. Flowers

± nodding with tepals held vertically with lowermost

slightly reflexed, zygomorphic with stamens unilateral,

bright to pale pink, occasionally white, yellow or white

edged in darker pink at tepal bases, lightly rose-scented

or unscented; perianth tube filiform, 2. 5^.0 nim long,

tightly clasping style; tepals ovate-elliptic, (8-) 10- 15 x

5-9 mm, inner 3 slightly larger than outer, lowermost

tepal held apart and usually somewhat reflexed. Stamens

unilateral; filaments inserted at mouth of tube, filiform,

1.6-3. 5 mm long, arching upward near tips, white to

pale yellow; anthers oblong-linear, 2. 3^.0 mm long,

horizontal or slightly pendent, thecae recurved and acute

at base, dehiscing incompletely from base by narrow

slits, yellow. Style dividing ± opposite base of filaments,

branches ± 1.5 mm long, pale yellow, falcate, tubular,

stigmatic apically. Capsules ± obovoid-oblong, 4-5 mm

long. Flowering time: mainly mid-Aug.-mid-Oct., occa-

sionally later. Figure 9.

Distribution and habitat: predominantly a species of

the west coast and near interior of Western Cape, Ixia

scillaris extends from the Cape Peninsula north to the

Gifberg and Kobee Mtns, along the coastal plain and

mountain ranges parallel to the coast as far inland as

the Cedarberg and the Pakhuis-Nardouw Mtns. Isolated

populations occur in the Spektakel Mtns in northern

Namaqualand, Northern Cape (Figure 10). Plants grow

FIGURE 9. — Ixia scillaris subsp. latifolia, Goldblatt & Manning 13482

(NBG). A, flowers; B, stamens and style branches, side view

with base of tepals; C, stamens and style branches, dorsal view.

Scale bar: 1 0 mm. Artist: John Manning.

FIGURE 10. — Distribution of Ixia scillaris subsp. latifolia, o.; subsp.

scillaris, •; subsp. toximontana. A; I. simulans, □; and I. tenuis,

A.

on a variety of soils but most commonly on heavy clays

derived from shale or granite. With so wide a range its

conservation status (LC), Least Concern, seems appro-

priate. The range of subsp. scillaris is, however, much

reduced and may be Vulnerable (VU).

Studies of the floral biology of Ixia scillaris (Gold-

blatt et al. 2000) show the flowers are adapted for buzz

pollination by anthophorine bees. Pollen remains in the

anthers until vibrated at high frequency by the wings

of visiting bees. Unusual for flowers with this pollina-

tion strategy, the pollen falls back into the centre of the

flower through narrow slits at the base of the anthers.

Like all species of sect. Dichone, the flowers do not pro-

duce nectar.

Diagnosis and relationships: the most common and

widespread species of sect. Dichone, Ixia scillaris is

recognized primarily by the nodding flower with verti-

cally oriented tepals with unilateral, slightly pendent

and often relatively long anthers, combined with a fan

of 3-7 basal leaves, (3)6-20 mm wide. Related I. eru-

bescens has narrow, ± falcate leaves, 2^(-6) mm wide,

with closely crisped margins, and is a smaller plant with

smaller flowers, the tepals 8-1 1 x 4_6 mm and anthers

2. 0-2. 5 mm long vs. tepals mostly 10-15 x 5-9 mm and

anthers 3^ mm long in I. scillaris. Apparently closely

allied Ixia simulans, with half to fully nodding flowers

and unilateral stamens with pendent anthers, has a sin-

gle basal, E linear foliage leaf and a second leaf sheath-

ing the stem for most of its length. Ixia tenuis from the

Piketberg is a particularly slender species with small

corms with finely fibrous tunics, just three, narrow, soft-

textured leaves and relatively small flowers with dark

magenta tepals. I. tenuis grows in a habitat unique for

species allied to /. scillaris, seeps on sandstone rocks,

growing in moss in soils less than 10 mm deep.

Variation: Typical Ixia scillaris from Darling and

Tulbagh to the Cape Peninsula has (4)5-7 relatively

narrow leaves, mostly 5-8 mm wide, with 3-5 second-

ary veins conspicuous only when dry, mostly branched

stems (rarely with up to 7 branches), and moderately

dense spikes with flowers spaced 8-12 mm apart. The

flowers are relatively small, with tepals 8-10 x 4—5 mm

and ± symmetrically arranged, although the lowermost

tepal is held slightly apart and recurved, and anthers

104

Bothalia 42,2 (2012)

2. 0-2. 8 mm long. In contrast, plants from the Olifants

River Valley and low elevations in the Cedarberg rang-

ing into the Bokkeveld and Spektakel Mtns are always

taller, with 2-A, broader leaves mostly 15-20 mm wide,

usually with at least two conspicuous secondary veins

in addition to the main vein, stems simple or with 1(2)

branches, and lax spikes with flowers distantly spaced,

15-25 mm apart. The flowers are larger, with tepals

14-16 X 5-9 mm, the upper tepals overlapping and

separated from the three lower tepals, and anthers 3.2-

4.0 mm long (Table 1). Although Lewis (1962) noted

the variation in leaf number and venation between the

southern and northern populations she preferred not to

recognize the two forms taxonomically as she believed

them to intergrade in leaf form and number to some

extent. She did not, however, note that the distinction

was accompanied by differences in size of the flowers,

which we consider as significant as the leaf differences.

A third set of divergent populations occurs on the Gif-

berg-Matsikamma Mtn complex. These consist of unu-

sually small plants, seldom exceeding 150 mm in height,

with small flowers (tepals ± 8 x ± 4 mm, anthers ± 2 mm

long), thus like those of the southern populations of Ixia

scillaris but with just three leaves (rarely two in depau-

perate individuals), recalling the northern populations, at

least in leaf number, although the leaves are shorter and

narrower.

Our taxonomic solution to the variation in Ixia scil-

laris is to recognize the three sets of populations as

separate subspecies, those of the larger-flowered, north-

ern populations as subsp. latifolia and the Gifberg-Mat-

sikamma populations as subsp. toximontana. We choose

subspecies rank because of the small overlap between

some populations of each subspecies and the compara-

tively small degree of difference between them relative

to that between species of sect. Dichone.

In addition to these major variants, Lewis recognized

populations with undulate leaves as var. subundulata.

The type of the variety is from east of Pakhuis Pass

{Barker 4727) and there are several more collections

from this area. Plants with undulate leaves also occur

locally near Tulbagh. Flowers of the Pakhuis-northem

Cedarberg have the relatively large flowers that cor-

respond with subsp. latifolia whereas those from the

south have smaller flowers of matching subsp. scillaris.

We see no value in recognizing undulate-leaved plants

taxonomically. Undulate leaf margins have evidently

evolved independently in different parts of the range of

Ixia scillaris.

History, the taxonomic history of Ixia scillaris

begins with a brief polynomial entry in A. van Royen’s

(1740) Florae leydensis prodromus, an account of the

plants grown in the botanical garden of Leiden Univer-

sity. Linnaeus (1753), in the first edition of the Species

plantarum, adopted van Royen’s species and, using his

actual phrase name, provided the specific epithet ramo-

sus to the description Gladiolus caiile ramoso,foliis lin-

earibus. The specimen is still at the Leiden Herbarium.

Then, in the second edition of his Species plantarum,

Linnaeus (1762) described Ixia scillaris, based on a

specimen in his own herbarium. Evidently Linnaeus

simply did not relate this specimen to the earlier name,

although he may well have seen Van Royen’s specimen

or the living plants when he was in The Netherlands

some 15 years earlier. Gladiolus ramosus was forgotten

for many years but was listed by Ker Gawler (1827) as

a synonym of Melasphaerula graminea (L.f.) Ker Gawl.

and by Lewis et al. (1962) as a synonym of the same

species under the name M ramosa (Burm.f) N.E.Br.

[the basionym for M ramosa is Phalangium ramosum

Burm.f (1768) and has no apparent connection with G.

ramosus L.]. F.W. Klatt (1894) cited G. ramosus L. as

the basionym for his combination Melasphaerula ram-

osa (L.) Klatt, an action that rendered Brown’s later

combination M ramosa (Burm.f) N.E.Br. a homonym.

Transfer of G. ramosus to Ixia is prevented by /. ram-

osa Ker Gawl., a replacement name for the homonym

I. scillaris Thunberg (1783), now Geissorhiza scillaris

A.Dietr. (Goldblatt 1985). Thunberg was uncertain of

the identity of 7. scillaris L., which he cited as a possible

synonym of 7 pentandra L.f

Key to subspecies

la Leaves (4—)5-7, lower two up to 10 mm wide, usually lan-

ceolate and lacking submarginal veins (margins rarely

TABLE 1 . — Significant taxonomic features of the subspecies of Ixia scillaris and allied I. tenuis. Observations are taken from living plants or from

well preserved specimens, bearing in mind that floral features may shrink up to 20% of original size, depending on the care with which

specimens are prepared.

Bothalia 42,2 (2012)

105

undulate); tepals 8-10 x 4-5 mm; anthers 2. 0-3. 5 mm

long subsp. scillaris

lb Leaves 2—4, lower two 3-25 mm wide, usually ± falcate,

sometimes with pair of prominent secondary veins,

these often submarginal (margins sometimes undulate to

crisped, then without prominent secondary veins); tepals

8-15 X 4—9 mm; anthers 2-4 mm long:

2a Leaves (9-) 12-25 mm wide; tepals 10-15 x 5_9 mm;

anthers mostly 3-4 mm long subsp. latifolia

2a Leaves 3-8 mm wide; tepals ±8x4 mm; anthers 1.6-2. 5

mm long subsp. toximontana

13a. subsp. scillaris

Gladiolus ramosus L.:37 (1753) [non Ixia ramosa

Ker GawL, (1802) = Geissorhiza scillaris A. Dietr.].

Melasphaerula ramosa (L.) Klatt: 203 (1894), epithet

misapplied to Melasphaerula. Type: South Africa, with-

out precise locality or collector, ex hort. Leiden, A. van

Royen s.n. (L: Herb. Royen, holo. — digital image!).

Ixia pentandra L.f.: 92 (1782). Agretta pentan-

dra (L.f.) Eckl. Type; South Africa, [Western Cape],

near Groene Kloof, without date, Thunberg s.n. (UPS-

THUNB, holo. — microfiche!).

Plants mostly 150-350 mm high, stem usually

branched, sometimes with up to 6 branches. Corm

mostly 12-14 mm diam. Leaves (4— )5-7, lanceolate to

sword-shaped or subfalcate, margins sometimes undu-

late, 5-8 mm wide, upper leaves progressively smaller

than lower. Spike mostly 10-1 5 -flowered, flowers

mostly 6-12 mm apart; bracts 3. 5^. 8 mm long. Flow-

ers pale to deep pink, darker at base of tepals, not pale in

throat, perianth tube 2. 5^.0 mm long, tepals subequal,

upper 5 tepals overlapping and evenly disposed with

lower laterals extended horizontally, lowermost held

apart, directed forward above base and then downward,

8-10 X 4-5 nun, iimer narrower than outer. Filaments

1. 5-2.0 mm long, anthers mostly 2-3 mm long. Flow-

ering time: mainly mid-Sept.-late Oct. (rarely in late

Aug.).

Distribution and habitat: restricted to the southwest-

ern Western Cape, subsp. scillaris extends from Tulbagh

and Darling to the Cape Peninsula and Somerset West

(Figure 10). The only record outside this area, a collec-

tion from Hawston near Bot River, Wilman s.n. NBG, is

the Clanwilliam form of Ixia scillaris, subsp. latifolia,

and we assume the label is incorrect. Subsp. scillaris is

conunon and fairly conspicuous where it occurs, mainly

on granite-derived sandy gravels.

Diagnosis: subsp. scillaris can be distinguished by

the several, (4)5-7, leaves mostly 5-8 mm wide, in

which only the main is prominent when alive (Table 1).

The flowers are slightly smaller than in subsp. latifo-

lia, with tepals 8-10 x 4—5 mm, the lowermost slightly

recurved and held separate from the upper five. The

anthers are also slightly smaller than in subsp. latifolia,

2-3 mm vs. mostly 3-4 mm long.

Representative specimens

WESTERN CAPE.— 3318 (Cape Town): Groenekloof [Mamre]

and hills nearby, (-AD), Oct., Ecklon & Zeyher Irid 241 (MO); Mamre

Hills (-AD), 26 Sept. 1941, Compton 11768 (MO, NBG); Cape Town,

above Camps Bay, (-CD), 26 Oct. 1944, Barker 3214 (MO, NBG);

Lions Head, (-CD), H. Bolus 2826 (BM, BOL, K); Paarl Mtn, near

Taal Monument, (-DA), 17 Oct. 1986, Goldblatt 7919 (MO, PRE);

Langverwacht above Kuilsrivier, sandy west-facing slope, (-DC), 24

Aug. 1973, Oliver 4362 (MO, NBG, PRE). 3319 (Worcester): fields

near Tulbagh, leaves undulate, (-AC), Sept. 1916, Marloth 7772

(PRE). 3418 (Simonstown): Helderberg, 240 m, (-BB), 17 Oct. 1948,

Parker 4360 (BOL, MO, NBG); Sir Lowry’s Pass, (-BB), Schlechter

5370 (BM, BOL, G, K); Vergelegen, Somerset West, (-BB), 13 Oct.

1982, Viviers 709 (PRE).

13b. subsp. latifolia Goldblatt & J.C. Manning, subsp.

nov.

TYPE. — Western Cape, 3218 (Clanwilliam): Cedar-

berg, Nieuwoudt Pass, burned slopes, (-BD), Goldblatt

& Porter 13475 (NBG, holo.; K, MO, PRE, iso.)

I. reflexa Andr.: t. 14 (1797). /. rotata [Ker Gawk]

in Andrews: 3 (1801), nom. illeg. superf. pro I. re-

flexa Andr. [Ker Gawk is the author of this publica-

tion (Stafleu & Cowan 1979), but internal evidence for

authorship is lacking]. Type: South Africa, without pre-

cise locality, cultivated in Great Britain by Messrs. Lee

& Kennedy and sent to them by J. Pringle but possibly

collected by F. Masson or W. Paterson (Gunn & Codd

1981), illustration in Andrews, The botanist’s repository

1: t. 14 (1797) — no preserved specimen known.

/. polystachya var. incarnata Andr.: t. 128 (1800).

Type: South Africa, without precise locality, cultivated

in Great Britain by Messrs. Lee & Kennedy and sent

to them by J. Pringle but possibly collected by Francis

Masson or William Paterson (Gunn & Codd 1981), illus-

tration in Andrews, The botanist’s repository 2: t. 128

(1800) — no preserved specimen known.

I. scillaris var. angustifolia Ker Gawk in Sims: sub

t. 542 (1801), nom. illegit. superf, including type of I.

polystachya var. incarnata Andr. Type: not designated,

two illustrations cited.

I. scillaris var. latifolia Ker Gawk in Sims: t. 542

(1801). Type: South Afiica, without precise locality,

cultivated in Great Britain, Masson s.n., illustration in

Curtis’s Botanical Magazine 15: t. 542 (1801) — no pre-

served specimen known.

I. scillaris var. subundulata G.J.Lewis: 177 (1962).

Type: Western Cape, Klipfonteiiu’and, 13 Sept. 1947,

Barker 4727 (NBG, holo.!, BOL!, iso.).

Plants mostly 300-500 mm high, stem simple or

with 1 or 2(3) branches. Corm mostly 14—20 mm diam.

Leaves 3 or 4, sometimes an additional leaf present at

base of a branch, lower 2 or 3 broadly falcate, (9-) 12-25

mm wide, usually with pair of conspicuous secondary

veins and often well developed submarginal vein, upper

leaf/leaves smaller, margins plane or undulate, then

without well developed secondary veins. Spike mostly

1 5-20-flowered, flowers mostly 14—20 mm apart; bracts

4— 5 mm long. Flowers pale to deep pink or white, then

flushed pale pink near tepal bases; perianth tube 3. 5-4.0

mm long; tepals subequal, upper 3 held close together,

lowermost held slightly apart arched backward, 10-15 x

5- 9 mm. Filaments 2. 0-3. 5 mm long, broader at base;

anthers 3-4 mm long. Style branches ± 2.5 mm long,

almost reaching anther bases. Flowering time: mainly

mid-Aug.-mid-Sept., rarely to early Oct. at higher eleva-

tions. Figure 9.

106

Bothalia 42,2 (2012)

Distribution and habitat: subsp. latifolia is centred

in the Olifants River valley and lower elevations of the

surrounding Cedarberg, Pakhuis-Bidouw Mts, extend-

ing from near Citrusdal to Bulshoek and locally in the

Bokkeveld Mtns. Isolated populations occur on the

Spektakel Mtns of northern Namaqualand, a notable

disjunction (Figure 10). It is replaced on the Gifberg/

Matsikamma Mtns by subsp. toximontana. Plants favour

rocky situations, mostly in sandy ground. Subsp. latifo-

lia is common throughout its range, but it flowers well

only after fire or clearing of the veld. The spring follow-

ing a fire plants are often seen in masses where before

the fire none were evident.

Diagnosis and variation: subsp. latifolia is distin-

guished by the presence of 3 or 4 leaves and by the rel-

atively larger flowers, the tepals 10-15 x 5-9 mm and

anthers 3^ mm long (vs. tepals 8-10 x 4-5 and anthers

2-3 m long). The upper three tepals are held more

closely together than the lower three and the lowermost

is conspicuously recurved.

Variation in leaf morphology in subsp. latifolia is

notable. Plants from Clanwilliam and nearby have a

conspicuous submarginal vein, but those from else-

where have a prominent secondary vein pair distant

from the margins, and plants from the Spektakel Mtns of

Namaqualand show no prominent secondary veins and

appear thinner in texture. No other feature seems to set

the Spektakel population apart. Populations east of the

Pakhuis-Nardouw Mtns have leaf margins slightly to

markedly undulate and without conspicuous veins apart

from the main vein.

Representative specimens

NORTHERN CAPE.— 2917 (Springbok): 14 miles [± 21 km) SW

of Springbok, (-DA), Acocks 19439 (NBG, PRE); Spektakel Pass,

E-facing slope near the top, (-DA), 26 Sept. 1974, Goldblatt 2804

(MO, PRE). Spektakel, (-DA), 25 Aug. 1941, Esterhiiysen 5853

(BOL). 3119 (Calvinia): between Nieuwoudtville and Oorlogskloof,

(-?AC), Aug. 1941, Leipoldt (BOL, PRE).

WESTERN CAPE.— 3218 (Clanwilliam): Zeekoe Vlei, (-BA), 19

Aug. 1896, Schlechter 8580 or 8500 (BM, G, K, MO, PRE); Clanwil-

liam, (-BB), 10 Sept. 1949, Steyn 516 (MO, NBG); Farm Kransvlei,

rocky sandstone ground, (-BB), 25 Aug. 2009, Goldblatt & Man-

ning 13482 (MO, NBG). 3219 (Wuppertal): Biedouw Valley on road

to Wuppertal, (-AA), 7 Sept. 1992, Goldblatt <6 Manning 9409 (MO,

NBG, PRE); Brandewyn River, (-AA), 29 Aug. 1941, Barker 1327

(BOL, NBG); Cedarberg, Langrug, (-AC), 12 Sept. 1982, Viviers 579

(PRE); Clanwilliam Dam to Algeria, (-AC), 9 Sept. 1976, Goldblatt

2567 (MO, PRE); banks of the Olifants River and at Villa Brakfontein,

(-BA), without date, Ecklon & Zeyher Irid. 243 or s.n. (G, MO, PRE,

SAM). Without precise locality: Grey’s Pass to Clanwilliam, Sept.

\94\, Leipoldt 3832 (BOL, PRE)

I3c. subsp. toximontana Goldblatt & J.C. Manning,

subsp. nov.

TYPE. — Western Cape, 3118 (Vanrhynsdorp): Gif-

berg, summit plateau in sandy ground over roeky pave-

ment, common after fire, (-DC), 1 Nov. 2011, Goldblatt

& Porter 13703 (NBG, holo.; K, MO, PRE, iso.).

Plants 120-250 mm high, simple or with 1 or 2,

rarely 3 branches. Corms globose-conic, mostly 15-18

mm at widest diam., tunics of hard, coarse fibres; bear-

ing conspicuous cormlets at base. Leaves (2)3(4), all ±

basal, mostly 3-8 mm wide, main vein prominent, sec-

ondary pair of veins hyaline when alive, raised when

dry, margins appearing thickened when dry. Spike

mostly 10- 15 -flowered, flowers mostly 8-12 mm apart;

bracts 3^ mm long. Flowers opening pale pink turning

deep pink next day, tepal bases yellow edged dark pink,

throat pale yellow, perianth tube ± 3 mm long, tepals

subequal, ±8x4 mm lowermost held slightly apart and

arched backward. Filaments 1.0- 1.6 mm long, anthers

1.6-2. 5 mm long. Style branches ±1.5 mm long, almost

reaching anther bases. Flowering time: late Sept.-mid-

Nov.

Distribution: restricted to the Gifberg and Mat-

sikamma Mtn complex of northern Western Cape, subsp.

toximontana grows in shallow, sandy ground overlying

sandstone pavement (Figure 10). The habitat is perma-

nently wet in the rainy winter and early spring months

but becomes dry by the end of September and remains

so though the summer and autumn. Most of its range is

virtually intact and pristine at present.

Diagnosis: subsp. toximontana is distinguished in

Ixia scillaris by its low stature, seldom exceeding 150

mm. combined with relatively small flowers and only 2

or 3 leaves, these short and relatively narrow, 3-8 mm

wide. The flowers have tepals ±8x4 mm and anthers

± 2.3 mm long, smaller than in subsp. latifolia or subsp.

scillaris (Table 1). In their small size they recall subsp.

scillaris rather than subsp. latifolia, in whose range it is

nested. The particularly small size of all parts may be an

adaptation to the habitat, nutrient-poor, shallow, sandy

ground.

Representative specimens

WESTERN CAPE. — 3118 (Vanrhynsdorp): Gifberg, (-DC), 17

Sept. 1961, Barker 9578 (NBG, PRE); top of Gifberg, (-DC), 23 Sept.

1962, Rycroft 2548 (NBG); summit of Gifberg Pass, open spaces and

shallow rock pans, (-DC), 9 Oct. 1973, Hall 4493 (NBG), 11 Oct.

1973, Bayliss 6153 (MO); Matsikammaberg, on recently burnt sand-

stone, (-DB), 11 Nov. 1985, Snijman 963 (NBG).

14. Ixia tenuis Goldblatt & J.C. Manning, sp. nov.

TYPE. — Western Cape, 3218 (Clanwilliam): Piket-

berg. Farm Noupoort, in wet mossy seeps on sandstone

rocks, (-DC), 2 Nov. 2011, Goldblatt & Porter 13710

(NBG, holo.; K, MO, PRE, iso.).

Plants slender, 180-350 mm high, stem simple or

rarely 1 -branched. Corms globose, 7-10 mm diam.,

tunics of fine fibres. Leaves mostly 3(2), occasionally 4

but then this very reduced in size, ± linear to narrowly

sword-shaped, 1/4-1/2 as long as stem, mostly 1. 5-3.0

mm wide, relatively soft-textured, pale grey-green,

when alive main vein slightly raised (appearing thick-

ened when dry), margins not raised (rarely undulate,

then blade up to 5 mm wide — Barker 4671, 5764, see

comments below). Spike mostly 4-10-flowered, flow-

ers mostly 10-15 mm apart; bracts 2-3 mm long. Flow-

ers deep magenta with tepal bases yellow edged with

darker magenta (or pale pink or white — Barker 4671,

5764), throat pale yellow, perianth tube 3. 0-3. 5 mm

long, expanded in upper 1 mm; tepals 8-9 x 4-5 mm,

oriented vertically but with lowermost tepals recurved

and ± horizontal. Stamens unilateral; filaments ± 2 mm

long; anthers 2. 0-2. 5 mm long. Style branches ± 1 mm

long, reaching to ± middle of filaments. Flowering time:

mid-Oct.-mid-Nov.

Bothalia 42,2 (2012)

107

Distribution and habitat: known from just a hand-

ful of sites, Ixia tenuis is restricted to rocky, sandstone

habitats in the Piketberg Mtns, mostly at elevations of

650-800 m (Figure 10). We collected the type popula-

tion on the Farm Noupoort on south-trending slopes of a

rocky hill growing in wet moss on shallow soil on sand-

stone outcrops. The remaining collections lack detailed

habitat notes. We note that the undulate-leaved variant

(see below) is from the foot of the Piketberg Range and

blooms earlier, in September. The species is rare, and at

present merits a conservation status of Vulnerable (VU),

but we see no imminent threat.

Diagnosis: Ixia tenuis is distinguished from the wide-

spread I. scillaris by its very slender habit, the stems

rarely branched, usually three, narrow leaves, mostly

1. 5-3.0 mm wide, and relatively small flowers with

tepals 8-9 x 4-5 mm and anthers 2. 0-2. 5 mm long

(Table 1). The corms are also relatively small, mostly

less than 10 mm in diameter and with relatively fine

tunic fibres. Corms of I. scillaris are larger, typically

12-18 (rarely only 10) mm in diameter, with relatively

coarse tunic fibres, and the species has broader leaves,

mostly 8-25 mm wide. We provisionally include collec-

tions by W.F. Barker {Barker 4671 and 5764) from De

Hoek and flowering in September here. These specimens

have slightly broader but shorter leaves than typical I.

tenuis and have undulate margins. This variant, which

we have not seen alive, was included by Lewis (1962)

as I. scillaris var. subundulata, the type of which we

include in /. scillaris subsp. latifolia.

Representative specimens

WESTERN CAPE. — 3218 (Clanwilliam): Piketberg Mtns, S side

of large boulders, 2 500 ft [800 m], (-DC), 3 Nov. 1951, Barker 7584

(NBG); Piketberg, top of mountain on Versfeld Pass, 650 m, (-DC),

4 Nov. 1988, Steiner 1863 (NBG); De Eloek, Piketberg, leaf margins

undulate, (-DC), 28 Sept. 1943, Barker 2560 (NBG), (flowers white),

12 Sept. 1947, Barker 4671 (NBG), 10 Sept. 1949, Barker 5764

(NBG).

15. Ixia flagellaris Goldblatt & J.C. Manning, sp.

nov.

TYPE. — [Western Cape]. 3219 (Wuppertal); Heu-

ningvlei, Groot Koupoort, common on wet sandy flats

burnt last summer, 3 150 ft [± 960 m], (-AA), 11 Oct.

1975, Kruger 1687 (NBG, holo.; PRE, iso.).

Plants 150-330 mm high, stem simple, rarely with

1 branch. Corm small, up to 10 mm diam., bearing sto-

lons up to 50 mm long from base, with a large termi-

nal cormlet up to 5 mm diam., tunics finely fibrous to

moderately coarse. Leaves 4, uppermost sheathing for

most of its length, foliage leaves with blades narrowly

sword-shaped to falcate, ± 1/3 as long as stem, mostly

2- 4 mm wide, main vein prominent, margins slightly

thickened. Spike mostly 5-9-flowered; bracts translucent

with purple veins, 2-4 mm long, outer 3-veined, acutely

3- toothed, inner slightly shorter than outer, 2-veined and

2-toothed. Flowers evidently zygomorphic, nodding

with tepals vertical, pink, throat and tepal bases yellow

edged with deep pink; perianth tube filiform, ± 3 mm

long; tepals ovate, ± 9.0 x 4.5 mm. Stamens evidently

unilateral; filaments slightly flattened, ± 2 mm long;

anthers oblong-orbicular, ± 3 mm long, yellow, thecae

recurved and acute at base, splitting incompletely from

base. Style dividing at base of filaments, branches ± 1.8

mm long, falcate, slender. Flowering time: Oct. and Nov.

Distribution and habitat: restricted to middle and

upper elevations in the Cedarberg, Ixia flagellaris is cur-

rently known from just two collections from the Cedar-

berg, one from Heuning Vlei in the north and the other

from Sneeuberg in the south (Eigure 8). The type col-

lection was made the season after a fire. The species

evidently grows on damp flats, most likely on peaty

sandstone ground, but this remains to be established. A

collection from Kobee Pass (see below) is provisionally

included here.

Diagnosis and relationships: the outstanding feature

of Ixia flagellaris is the corm, which is remarkable in

the /. scillaris complex in the finely fibrous corm tunics

and in the production of slender stolons (Latin: flagellae)

up to 50 mm long, each terminating in a relatively large

cormlet. No other member of sect. Dichone produces

stolons although they are known in a few species of

sect. Ixia and Morphixia (Goldblatt & Manning 2011).

1. flagellaris otherwise accords closely with I. scillaris

subsp. toximontana, notably in its relatively low stature,

150-330 mm high, and small flowers with tepals ± 9.0

X 4.5 mm, but differs in lacking prominent, thickened

leaf margins.. The late flowering and moist habitat, seeps

and marshy sites, are unusual for the complex, although

known in the Piketberg species, I. tenuis. Plants from

Kobee Pass {Hall 4517) cited below may belong here as

they have fine corm tunics and unthickened leaf margins

but they evidently do not produce stolons. Their exact

status remains uncertain.

Representative specimens

WESTERN CAPE. — 3219 (Wuppertal): Cedarberg, Sneeuberg,

Duiwelsgat Kloof, damp places, (-AC), 24 Nov. 1982, Viviers 789

(NBG). Provisionally included here: 3119 (Calvinia): summit of Kobee

Pass, open spaces in shallow pans, (-CA), 14 Oct. 1973, Hall 4517

(NBG).

16. Ixia simulans Goldblatt & J.C. Manning, sp. nov.

TYPE. — Western Cape, 3320 (Montagu): Langeberg

Mtns, gorge near Montagu hot springs above Keisie

River, on steep rocky sandstone slopes, (-CC), 1 Sept.

2007, Goldblatt & Porter 12946 (NBG, holo., K, MO,

PRE, iso.).

Plants 40-100 cm high, stems usually simple, rarely

with a single branch, sheathed basally by membranous

cataphylls. Corm ±10 mm diam., tunics of fine, netted

fibres. Leaves 2(3), lower 1(2) with narrowly sword-

shaped to linear blade with single prominent vein, ± 1/2

as long as stem, 1.5-6. 5 mm wide, margins thickened

and slightly raised, hyaline when dry; upper leaf sheath-

ing lower 1/2 of stem, free and unifacial distally. Spike

erect, fairly lax, weakly flexuose or straight, 4-8-flow-

ered; bracts membranous, translucent, purple at tips, ±

4 mm long, outer usually with 3 prominent veins and

3-toothed, inner 2-veined and forked at apex. Flowers

± nodding with tepals held vertically, zygomorphic with

stamens unilateral, bright pink (rarely pale pink), white

at tepal bases, unscented; perianth tube filiform, ± 4.5

mm long, tightly clasping style; tepals spreading, ovate-

elliptic, (10-)12-14 X 5-6 mm, inner whorl slightly

larger than outer. Stamens unilateral, filaments inserted

108

Bothalia 42,2 (2012)

at mouth of tube, filiform, ± 2.5 mm long, white, anthers

oblong, 2. 0-2. 5 mm long, yellow, horizontal or slightly

pendent, thecae acute and slightly recurved at base,

dehiscing incompletely from base by narrow slits. Style

dividing opposite lower 1/3 of filaments, branches ±

1.3 mm long, white, tubular, falcate, stigmatic apically.

Flowering time'. mid-Aug.-mid-Oct. Figure 11.

Distribution and habitat: Ixia simulans is currently

known from just four collections, three from the west-

ern Langeberg, between Dassieshoek near Robertson

and Swellendam, and one from Lemoenpoort in the

northern foothills of the Riviersonderend Mtns (Figure

10). Records are all from stony ground, and at the Kei-

sie River Gorge near Montagu hot springs plants grew

on steep, rocky, south-facing slopes. Plants are rooted

in shallow sandy soil, often growing in clumps of Res-

tionaceae. All known localities for I. simulans are in

virtually undisturbed montane vegetation and we see no

threat to the species: a conservation status of Least Con-

cern (LC) seems appropriate.

Diagnosis and relationships'. Ixia simulans is a sur-

prising novelty given that the western Langeberg has

been botanized for over two centuries and intensely so

in the past 50 years. The flowers closely resemble those

of I. scillaris in general appearance and have unilat-

eral, horizontal to slightly pendent anthers that dehisce

incompletely by narrow slits near the base. I. simulans

differs from I. scillaris most conspicuously in its sin-

gle, or rarely two, long, basal leaves with ± linear blades

1.0-4. 5 mm wide, spikes of only 4—8 flowers, relatively

short floral bracts ± 4 mm long and the spreading, sub-

equal tepals, those of the inner whorl being slightly

shorter than the outer. The tepals are inclined and all

held in the same plane whereas those of I. scillaris are

always held vertically and the lowermost tepal is weakly

to strongly recurved and held apart from the other five

tepals.

The range of Ixia simulans and I. scillaris comple-

ment one another: I. scillaris extends from the Cape

Peninsula and Somerset West in the south through the

Olifants River Valley, Cedarberg-Pakhuis Mtn complex

to the Bokkeveld Mtns, and locally well to the north in

the Spektakel Mtns west of Springbok whereas /. simu-

lans occurs well to the east of this line in the western

Langeberg.

Representative specimens

WESTERN CAPE. — 3319 (Worcester): Lemoenpoort, rocky

slopes, (-CD), 18 Aug. 1990, Bruyns sub Perry 3766 (NBG); Das-

sieshoek near Robertson, (-DB), 27 Aug. 2006, Goldblatt & Porter

12724 (MO, NBG, PRE). 3320 (Montagu): Langeberg Mtns, Marloth

Reserve ± 100 m from Glenstroom, 212 m, (-CD), 9 Oct. 2007, Turner

el at. s.n. (NBG).

17. Ixia collina Goldblatt & Snijman in South Afri-

can Journal of Botany 51: 68 (1985). Type: South

Africa, [Western Cape], Aan de Dooms, SW of Worces-

ter, Farm Alfalfa, Snijman 721 (NBG, holo!; MO!,

PRE!, SI, iso.).

Plants 500-900 mm high, stem sheathed below by

membranous cataphylls, usually 2-3-branched, branches

held at right angles to main axis, ascending distally,

subtended by short, acute bracts and prophylls. Corm

FIGURE 11. — Ixia simulans, Goldblatt & Porter 12946 (NBG). A,

flowering plant; B, outer (left) and inner (right) bract; C, half

flower; D, stamens and style branches, side view. Scale bar: 10

mm. Artist: John Manning.

Bothalia 42,2 (2012)

109

globose, 7-12 mm diam., tunics of medium-textured

fibres, often with cormlets at base. Leaves 3 or 4, lower

2 or 3 with sublinear to narrowly sword-shaped blades,

± 1/2 as long as stem, 4—8 mm wide, main vein mod-

erately thickened, uppermost leaf sheathing lower half

of stem. Spike flexuose, 7-16-flowered, lateral spikes

with fewer flowers; bracts membranous, translucent,

4. 5-7.0 mm long, outer with 3 prominent, purple veins,

3- or 5-toothed, inner 2-veined and 2-toothed. Flowers

zygomorphic, half nodding, pink, white to yellow edged

deep pink at tepal bases, sweetly scented, perianth tube

filiform, ± 5 mm long, expanded in upper 0.5 mm, tepals

subequal, ovate, ± obtuse, 9-12 x ± 5 mm. Stamens uni-

lateral, ± horizontal, inserted at mouth of filiform part of

tube, filaments ± 3 mm long, filiform below, curved at

right angles in upper 1 mm and becoming flattened and

as wide as anthers, anthers ± 3 mm long, oblong, yellow,

thecae acute and recurved near base, dehiscing incom-

pletely from base. Style dividing at base of filaments,

style branches ± 2.5 mm long, tubular, falcate, bifid at

apex, stigmatic apically. Flowering time: late-Aug.-mid

Sept.

Distribution and habitat. Ixia collina is a narrow

endemic of the mid Breede River Valley, its only known

locality the Farm Alfalfa near Aan de Dooms south of

Worcester (Figure 8). Plants grow on the southern slopes

of low clay hills above the Breede River flood plain and

adjacent to cultivated land. With so narrow a range Rai-

mondo et al. (2009) regard the species as endangered

(EN) and we agree. We recommend efforts be made to

locate additional populations but there is little doubt that

7. collina is extremely rare.

Diagnosis and relationships'. Ixia collina is easily dis-

tinguished from other members of ser. Dichone by the

unusual branching pattern: the relatively short branches

extend outward horizontally and curve upward distally

as well as by an unusual floral feature. The filaments are

unique in the section in being filiform below, curved at

right angles in the distal 1 mm and broadened so that

they are as wide as the anthers.

Additional specimen

WESTERN CAPE.— 3419 (Worcester): Aan de Dooms, SW of

Worcester, Scherpenheuwel, on Farm Reiersras, (-DC), Bruwer sub

Walters 750 (NBG).

EXCLUDED SPECIES

Ixia retusa Salisb., Prodromus stirpium in horto ad

Chapel Allerton vigentium: 35 (1796). The identity of this

species has remained a mystery as the description is inad-

equate to identify the plant (De Vos 1999) and no type

is known. Lewis (1962) suggested I. retusa might be 7.

stricta but with a query. Ker Gawler (in Sims 1801; 1803)

associated the name with what he called Ixia polystachya

(actually 7. scillaris, thus not the same as Linnaeus’s

species) [see extensive discussion under Taxonomic His-

tory]. Although we suspect that 7. retusa is a member of

sect. Dichone, it is impossible to tell which species and

the name must be rejected. Likewise, Klatt’s combina-

tion, Watsonia retusa (Salisb.) Klatt is rejected (see addi-

tional discussion under Ixia stricta).

ACKNOWLEDGEMENTS

Support for this study by grants 7316-02, 7799-05,

and 8248-07 from the National Geographic Society is

gratefully acknowledged. Collecting permits were pro-

vided by the Nature Conservation authorities of North-

ern Cape and Western Cape, South Africa. We thank

Ingrid Nanni, Elizabeth Parker, and Lendon Porter for

their assistance and companionship in the field; Roy

Gereau for advice with nomenclatural queries; Clare

Archer, South African National Herbarium, Pretoria

for help with several questions; Mary Stiffler, Research

Librarian, Missouri Botanical Garden, for providing

copies of needed literature; Sharon Bodine for help with

herbarium searches. We also thank the curators of the

following herbaria for allowing us access to their col-

lections or for the loan of material for extended study:

BOL, K, MO, NBG, PRE, and SAM (acronyms follow-

ing Holmgren et al. 1 990)

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Bothalia42,2: 111- 135 (2012)

Systematics of the hypervariable Moraea tripetala complex (Iridaceae:

Iridoideae) of the southern African winter rainfall zone

P. GOLDBLATT* and J.C. MANNING**

Keywords: Chromosome cytology, Iridaceae, Iridoideae, Moraea Mill., new species, southern Africa, taxonomy

ABSTRACT

Field and laboratory research has shown that the Moraea tripetala complex of western South Africa, traditionally treated

as a single species, sometimes with two additional varieties, has a pattern of morphological and cytological variation too

complex to be accommodated in a single species. Variation in floral structure, especially the shape of the inner tepals, degree

of union of the filaments, anther length and pollen colour form coherent patterns closely correlated with morphology of the

corm tunics, mode of vegetative reproduction, and in some instances capsule and seed shape and size. The morphological

patterns also correlate with geography, flowering time and sometimes habitat. It is especially significant that different vari-

ants of the complex may co-occur, each with overlapping or separate flowering times, a situation that conflicts with a single

species taxonomy. We propose recognizing nine species and three additional subspecies for plants currently assigned to M.

tripetala. M. grandis, from the western Karoo, has virtually free filaments and leaves often ± plane distally; closely allied

M. amabilis, also with ± free filaments and often hairy leaves, is centred in the western Karoo and Olifants River Valley. Its

range overlaps that of M. cuspidata, which has narrowly channelled, smooth leaves, linear inner tepals spreading distally and

filaments united for up to 1.5 mm. M decipiens from the Piketberg, M. hainebachiana, a local endemic of coastal limestone

fynbos in the Saldanha District, M. ogamana from seasonally wet lowlands, and early flowering M. mutila constitute the

remaining species of the complex in the southwestern Western Cape. M. helmei, a local endemic of middle elevations in the

Kamiesberg, Namaqualand, has small flowers with short, tricuspidate inner tepals. All but M. amabilis and M. mutila are

new species. We divide M. tripetala sensu stricto into three subspecies: widespread subsp. tripetala, subsp. violacea from

the interior Cape flora region, and late-flowering subsp. jacquiniana from the Cape Peninsula and surrounding mountains.

INTRODUCTION

The Afro-Eurasian and largely sub-Saharan genus

Moraea Mill. (Iridaeeae: Iridoideae) comprises ±215

species of cormous geophytes (Goldblatt & Manning

2009, 2010). Although florally diverse, the genus is rec-

ognized by a bifacial and channelled (rarely terete) leaf

blade (isobilateral leaves are ancestral for Iridoideae)

and a corm of a single intemode derived from a lateral

bud in the axil of the lowermost cataphyll. The major-

ity of Moraea species have Iris-like flowers with clawed

tepals, prominent, spreading outer tepal limbs marked

with nectar guides, and flattened, petaloid style branches

to which the anthers are appressed. Unlike Iris L., the

tepals of Moraea are normally free and the filaments

are at least partially united or rarely secondarily free in

a few species (always free in Iris). Species of Moraea

with other flower types (Goldblatt 1986b; 1998), usu-

ally with subequal tepals, reduced style branches and, in

some species, a perianth tube, are derived and limit the

utility of floral characters in circumscribing the genus.

Relatively widespread in the southern Africa win-

ter-rainfall zone, Moraea tripetala is one of the more

common species of subg. Vieusseuxia (D.Delaroche)

Goldblatt (± 30 spp.). The group is distinguished by the

flowers usually lasting three days (vs. fugaceous and

' B.A. Krukoff Curator of African Botany, Missouri Botanical Garden,

P. O. Box 299, St. Louis, Missouri 63166, USA. E-mail: peter.gold-

blatt@mobot.org

’* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, Claremont 7735, South Africa / Research Centre for

Plant Growth and Development, School of life Sciences, University of

KwaZulu-Natal, Pietermaritzburg, Private Bag XOl, Scottsville 3209,

South Africa. E-mail: J.Manning@sanbi.org.za.

MS. received: 2012-01-06.

lasting less than a single day, the plesiomorphic condi-

tion), a single, channelled foliage leaf, and inner tepals

variously modified in most species into trilobed or linear

structures, occasionally reduced to short hair-like cusps

or rarely absent (Goldblatt 1976a; 1986a). As currently

understood (Goldblatt 1976b) M. tripetala is recognized

by a blue, violet or purple perianth, reduced inner tepals

that are usually represented by short, hair-like cusps 2-3

mm long, filaments either free or shortly united for 1-2

mm (1/4 to 1/3 their entire length) and an associated

short style with the style branches appressed to the sta-

mens for almost their entire length. Variation in the col-

our and texture of the corm tunics, length and shape of

the leaves, in the size and shape of the inner tepals, and

the occasional development of pubescence on the stems

and leaves, was documented in some detail by Goldblatt

(1976b), who at that stage preferred not to accord any

of the variant forms taxonomic recognition. Goldblatt

(1976b) also noted that M. tripetala was variable cyto-

logically but the three populations sampled were diploid

with a basic chromosome number of x = 6, this shared

with all species of subgen. Vieusseuxia.

During extensive field work in western southern

Africa over the past decade we have encountered a wide

range of plants currently identified as Moraea tripetala.

The patterns of variation in the flower, especially the

form of the inner tepals, and associated differences in

corm, capsule and seed morphology, and in the means of

asexual reproduction have led us to the conclusion that

recognition of a single species for the complex does not

reflect biological reality. Particularly difficult to recon-

cile with a single species treatment for M. tripetala is

the co-occurrence of two morphological variants, either

flowering virtually side-by-side, or in bloom several

weeks or months apart in the same locality, and some-

112

Bothalia 42,2 (2012)

times differing in their habitat. With this in mind we

have undertaken an extensive review of the variation in

M thpetala. We conclude that a better treatment of the

complex, which we consider monophyletic, is to recog-

nize nine species, with M tripetala subdivided into three

subspecies.

MATERIALS AND METHODS

We first made field observations across the

entire range of the Moraea tripetala complex, not-

ing in addition to floral morphology, details of corm

tunics, mode of vegetative reproduction, and when-

ever possible capsule and seed morphology. These

observations were then integrated with an exami-

nation of specimens in herbaria with important

holdings of southern Africa flora, BOL, K, MO,

NBG, PRE, and S (abbreviations following Holm-

gren et al. 1990), plus a study of type material of the

named variants in the complex. Abbreviation of author

names follows Brummitt & Powell (1992).

Chromosome counts were determined from mitotic

squashes using root tips. Material for the original counts

reported here was prepared according to the protocol

described by Goldblatt & Takei (1993). The vouchers

are housed at the Missouri Botanical Garden Herbarium

(MO). Counts are based on samples of three to four indi-

viduals and, following standard practice in plant cytol-

ogy, are assumed to represent an entire population.

RESULTS

The lectotype of Moraea tripetala (designated by

Goldblatt 1976b), is a Thunberg collection from the

southwestern Cape, with filaments united for ± 1 mm,

inner tepals reduced to linear, acute cusps ± 3 mm long,

and a linear, narrowly channelled, glabrous leaf. Corms

are lacking in the type material but we are confident that

a range of plants recorded from Aurora on the west coast

of the Western Cape eastward to Knysna and mainly

flowering from August to the end of September accord

with the type. These often have slightly sweet-scented

flowers and the inner tepals vary from short hair-like

vestiges to linear structures up to 5 mm long. Apart

from plants matching the type relatively well, there is a

range of somewhat to significantly different populations

both in the southwestern Cape and in the interior, in the

western Karoo and Little Karoo. We describe the vari-

ant populations below, dealing first with plants from the

near southwestern Cape followed by those from the inte-

rior.

Variant 1: the first significant variant in the south-

western Cape is an early flowering taxon, mostly bloom-

ing from mid-August to early September, often with

relatively broad leaves. These are sometimes plane dis-

tally, 3-5(-7) mm wide and laxly twisted, ± half as long

to ± as long as the stem, or rarely exceeding it. Plants

occur at low elevations on clay or loamy soils. The inner

tepals are linear to narrowly lanceolate, up to 12.5 mm

long, and differentiated into an erect portion (equivalent

to the claw of the outer tepals) and an outspread portion

(equivalent to the limb) that is either slightly expanded

at the base or, in some populations (e.g. Goldblatt 2310

MO), with two obtuse, lateral lobes, rendering the limb

± trilobed. The filaments are united for 1.0-1. 5 mm,

rarely only ± 0.5 mm. In two collections the leaves and

stem of some plants are thinly hairy (e.g. Nordenstam

& Lundgren 1998 MO, NBG, S; Acocks 1912 S). This

variant was first recognized at species rank by C.F. Eck-

lon (1827) as Vieusseuxia mutila and later by J.G. Baker

(1904) as Moraea punctata but has remained poorly

understood. Moraea monophylla (Baker 1906) and M.

tripetala var. mutila (Baker 1896) represent the same

variant.

Plants from the immediate area of Sir Lowry’s Pass,

east of Cape Town, flowering in August, have inner

tepals =E 10 mm long, consisting of a limb with a short

central tapering cusp and short rounded lateral lobes.

They are somewhat out of range for variant 1 and have a

narrow, channelled, smooth leaf (e.g. Loubser 872 NBG;

Goldblatt 2506 MO). These plants may belong here or

alternatively may be hybrids with M unguiculata or

another species: we discuss them below in more detail.

Variant 2\ a second southwestern Cape variant is the

plant G.J. Lewis (1941) named Moraea tripetala var.

jacquiniana (= M. jacquiniana Schltr. ined.). Often

shorter in stature, 140-300 mm high and with smaller

flowers than in typical M. tripetala, it usually has excep-

tionally slender leaves, mostly 2-3 mm wide, V-shaped

in cross section. When dry the blades are closely folded

together and appear terete. Flowering in this plant is

from mid-November to January on the Cape Peninsula

(Lewis 1950), but sometimes in late October elsewhere.

Illustrated in Flora of the Cape Peninsula (Maytham

Kidd 1950), the flowers are often dark purple, but some-

times light mauve to ± blue as in M tripetala sensu

stricto, and as far as known, consistently have white

nectar guides. The linear inner tepals are 5-6 mm long

(e.g. Pillans 10279 BOL, MO), thus slightly more than

1/2 as long as the outer tepal claws and reaching to

between the base and middle of the anthers. The outer

tepals have limbs 11-16 mm long and claws 8-11 mm

long (vs. limbs 12-18 and claws 10-12 mm long in M.

tripetala sensu stricto). At several localities, notably in

the southern Cape Peninsula, at Jonkershoek and in the

Grabouw area, both typical M. tripetala and var. jac-

quiniana occur, the former blooming two to four months

earlier, late July to September. M. pulchra Eckl. (1827)

from ‘Hottentotshollandkloof’ collected in flower on 25

November represents the earliest name for this plant at

species rank.

Variant 3: restricted to lowland wetlands of the south-

western Cape and flowering in September, this vari-

ant has an unusual, narrow ovary, cylindric rather than

ovoid, 10-13 mm long, and darkly lined vertically on

locules, and nearly cylindric capsules (15-)20-24 mm

long, somewhat angled on the locules. Typical M. trip-

etala has an ellipsoid ovary 6-10 mm long and capsules

8-14 mm long, both round in cross section. The flow-

ers of this variant are also unusual in having the outer

tepal limbs pale blue with darker veins radiating from a

yellow nectar guide. Plants are always relatively short,

rarely exceeding 200 mm, have leaves usually shorter

than the stems, and small corms with blackish, wiry

tunics. There is also invariably a small cormlet in the

axil of the foliage leaf, and plants often grow in small

Bothalia42,2 (2012)

113

groups, the result of vegetative reproduction from these

cormlets. Typical M tripetala does not normally pro-

duce cormlets either in the leaf axils or at the corm base,

and is always a solitary plant, favouring well-drained

habitats. Variant 3 extends from Voelvlei in the north to

Harmony Flats at Strand in the south.

Variant 4\ a last variant in the southwestern Cape

is a vegetative apomict, restricted to limestone pave-

ment or calcareous sands in the Saldanha Bay area, and

blooming in August and early September, rarely later.

Relatively short, up to 300 mm high, plants have lilac or

pale to mid-blue flowers with pale yellow nectar guides,

outer tepals 27-29 mm long and unusual inner tepals ±

4 mm long, spindle-shaped, sometimes oblanceolate and

obscurely 3-lobed, tapering distally with the apex curv-

ing inward, sometimes the inner distal surface pilose.

Unusual for the complex, it has pale grey-blue anthers

and off-white or yellow pollen. Capsules never develop

to maturity and are shed several days after flowers fade

and seeds are not formed. Microscopic examination of

pollen confirms that the grains are all malformed. Propa-

gation of new plants is accomplished by the production

of several cormlets bom in the axil of the foliage leaf

and sometimes the lowermost sheathing leaf Non-flow-

ering individuals produce similar cormlets at the base of

the main corm. This variant co-occurs with typical M

tripetala, which blooms in the same sites in late Sep-

tember and early October, and has paler blue flowers and

dark blue or purple anthers and red pollen typical of the

species.

Variant 5: some plants from the Piketberg assigned to

Moraea tripetala by Goldblatt (1976b) have large inner

tepals 7-10 mm long with distally expanded limbs that

are ± trilobed and tapering to an attenuate, twisted tip.

These plants are also unusually tall, reaching up to 450

mm, but the flowers are relatively small for the com-

plex, with outer tepals 20-23 mm long and claws 8-9

mm long. The filaments, only ± 4 mm long, are united

for ± 2 mm. In general appearance these plants resem-

ble M unguiculata Ker Gawl, but the filaments, united

for less than 1/2 (slightly more than 1/3) their length, are

characteristic of the M tripetala complex. Both typical

M tripetala and white-flowered M. unguiculata occur

locally in the Piketberg, where the former has the short,

hair-like inner tepals typical of the species.

Variant 6: outside the immediate southwestern Cape,

in the western Karoo, plants included in Moraea trip-

etala usually have free filaments (sometimes united

basally for < 0.4 mm), inner tepals reduced to small

hair-like cusps mostly 1-2 mm long, a linear leaf, shal-

lowly channelled below but often plane distally, and

light brown corm tunics of medium-textured to soft

fibres. Plants always form clonal colonies, the result of

vegetative propagation by the production of two or more

cormlets at the base of the flowering stem that replace

the parent corm, and/or by axillary cormlets (Figure 14).

Populations of this variant fall into two main groups.

The first of these, from the western Karoo and the Oli-

fants River Valley and surrounding mountains, mostly

from lighter, sandstone or shale-derived soils, has

smaller flowers with outer tepals 23-25 mm long (claws

10-14 mm), anthers 4.5-7.0(-8.0) mm long and ±

as long as to 1.5 x as long as the filaments, and either

white to yellow pollen (eastern populations) or red pol-

len (western populations) (Figure 14). Pollen colour is

lost in older collections but our observations are based

on examination of living plants across almost the entire

range of this morph. The 5(6)-sided seeds are 1. 7-2.0

X ± 1 mm and stand out in having pale, spongy, ridged

angles and an extended pale spongy mass at the micro-

pylar end. Seeds of typical M. tripetala and related

variants are 1.0-1. 3 x ± i mm, with sharply raised, pale

angles and a finely wrinkled surface through which the

underlying dark brown seed body is evident. This vari-

ant matches the type of M. amabilis Diels.

Plants closely resembling this variant growing on

sandstone pavement in the Bokkeveld and Gifberg-Mat-

sikamma Mtns are often particularly short, usually < 1 80

mm high. Capsules and seeds in these plants are consist-

ent with this variant.

A second western Karoo variant comprises large-flow-

ered plants, mostly of dolerite-derived soils, with outer

tepals 30^0 mm long (claws 12-18 mm long), anthers

8-11 mm long and ± twice as long as the filaments, and

consistently red pollen (Figure 15). The capsules are

16-19 mm long, narrowly ellipsoid-oblong with a mark-

edly thickened apical rim. Seeds in the few fruiting col-

lections that we have been able to examine have excep-

tionally large, brown seeds, 2-3 x 1.8-2. 3 mm.

Variant 7: also in the western Karoo and adjacent

interior southern Cape, another variant of the complex

has linear inner tepals (8-)10-15 mm long, ascending

below and spreading distally, filaments united for 0.5-

1.8 mm, and narrowly charmelled leaves with the leaf

halves often appressed. The relatively large corms have

tunics of wiry, usually thickened fibres, and cataphylls

that decay into a particularly prominent collar of verti-

cal fibres around the base. The filaments are (3-)4-6

mm long and the anthers, 4-8 mm long, with red pollen.

Late flowering, these populations seldom bloom before

the last week of September and only in late October at

higher elevations in the Swartberg Mtns. Plants are

sympatric in the Roggeveld and Klein Roggeveld with

the western Karoo morph of variant 6, which has short,

hair-like inner tepals and pale corm tunics, and their

flowering times often overlap (e.g. Goldblatt & Porter

13461 and 13462, MO, NBG, from the Farms Fortuin

and Nuwerus). The few plants we have seen with mature

capsules have distinctive seeds with a spongy testa

slightly thicker on the angles.

Variant 8: plants from the Kamiesberg, Namaqua-

land, discovered only in 2009 by Cape Town botanist

N.A. Helme, and evidently belonging to the Moraea tri-

petala complex, have filaments united in the lower 1.5

mm but differ notably in their short, trifid inner tepals,

represent the last significant variant of the complex (Fig-

ure 4). Apart from the unique inner tepals, the flowers

are small, with outer tepals 21.0-23.5 mm long bear-

ing bright yellow, velvety nectar guides. The outer tepal

claws have a pair of marginal teeth just below their api-

ces.

Even with the exclusion of these variants, the remain-

ing populations of Moraea tripetala are still variable.

114

and we discuss the major patterns below under that spe-

cies. Extending from Aurora and the Piketberg on the

west coast to the Cape Peninsula and thence across the

southern Cape to Knysna, plants have moderately-sized,

pale blue, purple or violet flowers with white or yellow

nectar guides (many populations are variable for flower

colour), inner tepals hair-like and rarely > 4 mm long,

and the ovary is ellipsoid and 8-10 mm long. Vegeta-

tive reproduction via axillary or basal cormlets is rare.

Among these populations, plants from the southern Cape

sometimes lack inner tepals entirely and those from the

Langeberg foothills may have free filaments. Neither

trend is consistent as far as we can determine. From the

few fruiting specimens available, seeds of what we con-

sider typical M. tripetala are consistently small, with

pale, raised, ± winged angles.

DISCUSSION

Species, subspecies or merely ecotypes?', the immedi-

ate question our observations raise is whether or not any

of the several morphological variants should be recog-

nized taxonomically, thereby reducing the otherwise flo-

rally distinctive Moraea tripetala to just one of several

taxa that are only moderately distinctive florally and not

easily distinguished without examination of corms, cap-

sules, and seeds. We conclude that the patterns of vari-

ation that we now recognize renders the single species

solution unacceptable, particularly because of the cor-

related character differences in the conn, leaf, flower,

and seeds, and the examples of sympatry of two vari-

ants at some sites, either flowering at the same time or

flowering weeks to months apart. Moreover, the range

of habitats and flowering times would be remark-

able for a single species and unacceptable for the lack

of conformity with any but the broadest morphological

species concepts. Similarly, treatment of the major vari-

ants as subspecies violates any biological species con-

cept. We conclude that all the distinctive entities that

co-occur and have overlapping flowering times should

be treated as separate species. Likewise, co-occurring

variants that have flowering times separated by weeks

or months require taxonomic recognition if they can be

readily identified by more than one unique morphologi-

cal marker.

This framework has been the underlying philosophy

in our decision to dismember Moraea tripetala sensu

Goldblatt (1976b). We also note that Baker (1896) in

Flora capensis recognized one variety in the species and

later added two more (Baker 1904, 1906). Lewis (1950)

recognized M. tripetala as one species with three vari-

eties in Flora of the Cape Peninsula alone. Three of

the eight segregate species we recognize already have

names in the literature, providing a precedent for our

revised taxonomy [M amahilis, M. tripetala var. jac-

quiniana (= Vieusseuxia pulchra) (Variant 2), M. punc-

tata (= Vieussuexia mutila = M. tripetala var. mutila)

(Variant 1)]. To these we now add M. decipiens (variant

5); M. hainehachiana (variant 4); M. ogamana (variant

3); M. cuspidata (variant 7); and M. helmei (variant 8).

We also recognize M. amahilis and M. grandis from the

western Karoo for the two subsets of variant 6.

With the recognition of nine species in the Moraea

tripetala complex, section Vieusseuxia now includes 36

Bothalia 42,2 (2012)

species, and the genus Moraea ±215 species (Goldblatt

& Manning 2010 and unpublished).

Floral and reproductive biology, as might be

expected in species with similar flowers, all members

of the complex share the same pollination system, large

bodied, mostly anthophorine bees (usually Amegilla and

Anthophora spp.), or sometimes Apis mellifera (Gold-

blatt et al. 2005). Our impression is that the species of

the complex are remarkably successful and it is rare to

see plants later in the season without multiple capsules

with full complements of seeds; autogamous selfing

can be dismissed because in these iris-type flowers the

anthers do not reach the stigmatic lobes (Goldblatt 1998)

and we infer that insect mediated pollen transfer is nec-

essary to achieve pollination.

Propagation by seed is complemented in several spe-

cies either by production of cormlets in the leaf or cata-

phyll axils and at the base of the corm {Moraea haine-

bachiana, M. ogamana) or sometimes by production

of two new corms in place of the parent corm {Moraea

amahilis and often M grandis). Enhanced vegetative

reproduction in these species results in plants form-

ing clonal colonies. These two vegetative propagation

strategies are uncommon elsewhere in the M. tripetala

complex. The lowland wetland populations we treat as

M. ogamana consistently produce a single cormlet in the

foliage leaf axil but the vegetative apomict, M. haine-

bachiana produces multiple cormlets in the foliage leaf

axil as well as at the corm base, a feature particularly

well-developed in non-flowering individuals. This last

species produces malformed pollen, and the capsules

are invariably shed soon after flowers wilt, without the

ovules developing into seeds.

Chromosome cytology, as noted by Goldblatt (1976a),

chromosomal karyotypes are variable for populations

then included in Moraea tripetala. Sampling is limited

(Table 1) but we note the following. M amahilis, M.

cuspidate, and M grandis have the longest three chro-

mosome pairs either metacentric or sub-metacentric

(Figures lA, B) and a small satellite on the fourth long-

est ± acrocentric pair (Goldblatt 1971 and Table 1, as sat

type 1). This karyotype is the common one in subgen.

Vieusseuxia. The karyotype of M. mutila has small sat-

ellites on the two longest, submetacentric chromosome

pairs (sat type 3) (Figure 1C). In contrast, plants of M.

tripetala subsp. tripetala and subsp. violacea (Table 1),

have very large satellites on a long, telocentric chromo-

some pair (sat type 2). In three of four populations of

subsp. tripetala sampled a second large satellite (Figure

ID) is evident on a relatively short acrocentric chromo-

some pair (sat type 2a) (poor preparation in the fourth

sample makes it uncertain whether the second pair of

satellites is present or not). These examples, inadequate

though they are, show that chromosomal rearrangements

via inversion or translocation occur with some frequency

in the complex and potentially provide a genetic basis

for reproductive isolation because of meiotie disruption

in any inter-populational hybrids. The three karyotypes

also provide support of our revised taxonomy in show-

ing different karyotypes, consistent within species. M.

decipiens, M. hainehachiana, M. helmei, M. ogamana,

and M. tripetala subsp. jacquiniana remain to be exam-

ined cytologieally.

Bothalia 42,2 (2012)

115

TABLE 1. — Chromosome counts in Moraea tripetala complex with corrected taxonomy. Satellite characteristics are as follows: small on acrocen-

tric chromosome, sat type 1; large on telocentric chromosome, sat type 2, type 2a with second large satellite on short acrocentric pair; small

on large submetacentric chromosome, sat type 3. Original counts are in bold type in column 2 and their voucher specimens are at the Missouri

Botanical Garden Herbarium (MO).

TAXONOMY

Key to the species of the Moraea tripetala complex

la Inner tepals 7-20 mm long, linear or expanded and lobed in

middle or 3-lobed with central lobe tapering and twisted:

2a. Inner tepals 7-9 mm long, 3-lobed with longer, tapering,

often twisted central lobe; style crests ± 5 mm long;

plants from middle elevations of the Piketberg, flower-

ing mainly November 1 . M decipiens

2b. Inner tepals 7.5-15.0 mm long, linear throughout or

expanded and lobed in middle; style crests 5-10 mm

long; plants widespread on a variety of soils, flowering

mainly August to October:

3a Foliage leaf usually shorter than stem, even half as long,

leaf blade distally at most shallowly channelled or ±

plane and laxly twisted, mostly 3. 5-5.0 mm wide; leaf

blades and/or stem or spathes and sheathing leaves

sometimes hairy; inner tepals 7.5-12.5 mm long, often

expanded in middle, sometimes into small rounded

lobes at base of spreading distal half; ovary 7-8 mm

long; plants flowering mainly in August) September)

3. M mutila

3b Foliage leaf as long as or more often exceeding stem,

sometimes twice as long; leaf blade narrowly chan-

nelled with leaf halves often folded together, 1. 5-3.0

mm wide; vegetative parts never hairy; inner tepals

8-15 mm long, linear for entire length, spreading in

distal half to one third; ovary 11-15 mm long 4. M. cuspidata

lb Inner tepals (l-)2-6 mm long or occasionally absent, either

3-lobed in upper third or reduced to hair-like or lanceolate

to linear cusps, acute and usually widest at base:

4a Inner tepal ± 4 mm long, 3-lobed in upper third; filament

column and lower abaxial surface of inner tepals puberu-

lous; plants of the Kamiesberg, Namaqualand . . . 2. M. helmei

4b Inner tepal variable in length (1-6 mm), a hair-like, linear

cusp; filament column and lower abaxial surface of inner

tepals smooth; plants of the western and southern Cape

and western Karoo:

5a Axil of foliage leaf bearing several dark-coloured corm-

lets breaking through leaf sheath; plants never forming

mature capsules or seeds; anthers pale blue and pollen

blue or white to yellow; plants restricted to Western

Cape west coast near Saldanha on limestone pavement

or calcareous sands 6. M hainebachiana

5b Axil of foliage leaf rarely with a solitary cormlet

enclosed in sheath or without cormlets; anthers and

pollen orange-red or white (rarely yellow); plants

always forming mature capsules and fertile seeds;

plants widespread from the Western Cape coast and

Western Karoo on various soils:

6a Filaments usually united for 0.5-1. 5 mm (rarely free);

plants not forming colonies, occasionally a sin-

gle cormlet present in axil of foliage leaf; leaves

narrowly channeled and leaf halves often closely

appressed; stems and abaxial leaf surface smooth:

7a Ovary narrowly ellipsoid, mostly 4—9 mm long;

capsules ellipsoid, 8-14 mm long; foliage leaf (or

leaves) usually as long as or longer than stem; axil-

lary cormlets absent; pollen usually orange-red,

occasionally white 5. M tripetala

7b Ovary linear-cylindric, 10-13 mm long; capsules ±

cylindric (15-)20-24 mm long; foliage leaf often

shorter than stem, sometimes half as long; bearing

a single cormlet in axil of foliage leaf; pollen white

7. M ogamana

6b Filaments ± free or united basally for up to 0.4 mm;

plants usually forming colonies due to vegetative

reproduction by axillary cormlets and/or two new

cormlets replacing parent corm; leaves widely chan-

neled, often ± plane distally; plants sometimes vel-

vety on stems and abaxial surface of leaves:

8a Flowers with outer tepal limbs (9-)ll-18 x 10-14

mm, always as long as or longer than wide; anthers

4.5-7.0(-8.0) mm long and up to one and a half

times as long as filaments; corm tunics usually of

hard, coarse, light brown (occasionally dark brown)

fibres 8. M amabilis

8b Flowers with outer tepal limbs 15-20(-23) x 15-20

mm, often ± as long as wide; anthers 8-11 mm

long and ± twice as long as filaments; corm tunics

usually of soft, fine (rarely medium-textured), light

to dark brown fibres 9. M grandis

1. Moraea decipiens Goldblatt & J.C. Manning, sp.

nov.

TYPE. — Western Cape, 3218 (Clanwilliam): Piket-

berg, Farm Noupoort, south-trending, rocky sandstone

slope in shallow pockets of soil, (-DA), 2 Nov. 2011,

Goldblatt & Porter 13709 (NBG, holo.; K, MO, PRE,

iso.).

Plants 300^50 mm high. Corm 9-12 mm diam.

with pale, fibrous tunics. Stem usually simple, rarely

1 -branched, glabrous. Foliage leaf solitary, linear, con-

duplicate with margins tightly folded together, 0.7-1. 5

116

Bothalia 42,2 (2012)

° i()( li'ii ii«

FIGURE 1. — A. Mitotic metaphase and idiogram of karyotype of

Moraea grandis, Goldblatt lOJ); B-D, karyotypes from mitotic

metaphases with chromosomes arranged in pairs for comparison;

B. M amabilis, Goldblatt 3852; C. M. mutila, Goldblatt 6127). D.

M. tripetala subsp. tripetala, Goldblatt 2607B).

mm diam., exceeding stem and often trailing above,

sometimes with axillary cormlet; sheathing cauline

leaves 45-55 mm long, green with dry attenuate apices.

Rhipidia mostly 3-5 -flowered; spathes green with dry,

brown, attenuate tips, inner 35^0 mm long, outer ± half

as long as inner. Flowers pale to deep purple, outer tepal

limb bases with white, wedge- or V-shaped, minutely

papillate nectar guides dark purple in centre; outer

tepals 20-23 mm long, claw 8-9 mm long, with wide,

dark violet median stripe, limb 12-15 x 6-8 mm; inner

tepals 7-10 long, violet, claw suberect, plane, expanded

to 2 mm wide at apex, often into rounded lobes, limb

3-4 mm long, tapering to attenuate tip, laxly twisted dis-

tally. Stamens with filaments 3. 5^.0 mm long, united

in lower ± 2 mm; anthers 4. 5-6.0 mm long, reaching or

shortly exceeding stigma lobe, dark purple-black; pollen

orange-red. Ovary narrowly oblong-truncate, 7-9 mm

long, usually exserted when flower mature; style ± 2 mm

long, dividing ± 0.5 mm above top of filament column,

branches, 7-9 x 1-2 mm; stigma shallowly bilobed;

style crests narrowly wedge-shaped, ± 5 mm long, erect.

Capsules narrowly ovoid, 8-11 mm long. Seeds 5(6)-

sided, 1.2-1. 5 mm x ± 1 mm, facets slightly wrinkled,

brown seed body visible though testa, angles between

facets forming narrow, raised, pale, slightly spongy

ridges. Chromosome number unknown. Flowering time;

late Oct.-mid-Nov. Figure 2.

Distribution; known only from the western half of the

Piketberg in Western Cape (Figure 3), Moraea deeipiens

is evidently restricted to middle elevations of the range,

rather than the emergent peaks, and based on our own

collection, occurs on stony sandstone slopes in shallow

pockets of soil among Restionaceae and low shrubs.

None of the other three collections that we have seen

include information about the habitat.

Diagnosis; named for the deceptive {deeipiens =

Latin deceiving) appearance of the flowers to those of

M. tripetala, M. deeipiens, nevertheless, differs in hav-

ing the inner tepal claws distinctly expanded distally,

sometimes into rounded lobes, with the limb linear-

attenuate and curved inward above. The inner tepals

recall those of 'M. unguiculata and M. algoensis but

unlike those species, the filaments of M. deeipiens are

only shortly united for ± 2 mm and the style branches

diverge above the fused part of the filaments whereas in

M. unguiculata and M algoensis the filaments are united

almost to their apices in a relatively thick column. The

style branches of M deeipiens are relatively short, ± 7

mm long, unlike those in M. tripetala which are usually

9-12 mm long. The markings on the outer tepals are also

notable, consisting of a wedge- or V-shaped white zone

surrounding a dark purple basal mark that is continuous

as a wide, longitudinal purple streak on the claw. Plants

from Pakhuis Pass, flowering in late October {Flelme

5726, NBG) bear a superficial resemblance to M. decipi-

ens in size of the flowers, colour of the nectar guides and

the broad inner tepals but the filaments are united for

2.5 mm and 6.5 mm long, thus longer than the anthers,

which are just 4 mm long. More material of this plant is

needed before an informed decision can be made about

its status but we suspect it represents yet one more unde-

scribed species of subgen. Vieusseuxia.

Typical Moraea tripetala itself also occurs at higher

elevations on the Piketberg, e.g. Linder 414 BOL, from

Levant Mtn where M. deeipiens also occurs, but flowers

some two to four weeks later. Early flowering M. mutila

of the complex occurs in renosterveld on the lower east-

ern slopes of the range and has inner tepals broadly

similar to those of M. deeipiens, although narrower and

longer. The population at the type locality was relatively

large and plants showed no variation in critical features,

thus unlikely to be hybrid between M. tripetala and M.

mutila. Barker’s collection of M. deeipiens likewise

comprises several plants {Barker 7563), all uniform for

flower features, and the pollen appears normal under the

microscope.

Additional specimens

WESTERN CAPE.— 3218 (Clanwilliam): hills NW of Mouton’s

Vlei, (-DA), Nov. 1934, Pillans 7489 (BOL); Piketberg, Levant Mtn,

Farm Avontuur, (-DA), 5 Nov. 1973, Linder 81 (BOL); Piketberg,

without precise locality, 3 Nov. 1951, Barker 7563 (NBG).

2. Moraea helmei Goldblatt & J.C. Manning, sp. nov.

TYPE. — Northern Cape, 3018 (Kamiesberg): south-

ern Kamiesberg south of Farm Karas, near top of Lang-

kloof, 1 125 m, (-AB), 28 Oct. 2011, Goldblatt & Por-

ter 13688 (NBG, holo.; MO, iso.).

Plants 250-350 mm high. Corm 9-12 mm diam.

with pale, fibrous tunics. Stem simple or with 1 or 2

Bothalia 42,2 (2012)

117

FIGURE l.—Moraea decipiens, Goldblatt & Porter 13709. A, flowering stems and corm; B, inner tepal variation; C, inner perianth whorl plus fila-

ments and style; D. capsules. Scale bar: A, D, 10 mm; B, C, 2 mm. Artist: John Maiming.

118

Bothalia 42,2 (2012)

FIGURE 3. — Distribution of Moraea cuspidala, o; M. decipiem, •; M

helmet. A; and M mutila, A .

short branches, glabrous. Foliage leaf solitary, lin-

ear, channelled, remaining ± erect, ± 2 mm wide when

opened flat; sheathing cauline leaves 2 or 3, 35^5

mm long, green, attenuate, becoming dry at tips. Rhip-

idia 2(3)-flowered; spathes green with dry tips, attenu-

ate, inner 40^5 mm long, outer ± two thirds as long,

entirely sheathing. Flowers pale blue to violet, outer

tepal limbs darkly veined, with narrowly triangular,

shortly velvety, yellow nectar guides at base of outer

tepal limbs outlined pale blue to dark violet; outer tepals

21.0-23.5 mm long, claw 8-10 mm long, with a shallow

raised tooth either side just below apex, limb 12.5-13.5

X 7-8 mm; inner tepals short, ± 4 mm long, 3-forked in

upper 1 mm, inner lobe slightly larger than laterals, ±

puberulous. Stamens with filaments ± 4 mm long, united

in lower 1.5 mm, column puberulous; anthers ±6.5 mm

long, dark violet; pollen yellow. Ovary narrowly ellip-

soid, ± 7 mm long, initially partly included, ultimately

fully exserted; style ± 2 mm long, dividing at top of fila-

ment column, branches linear, 9x1 mm; style crests ± 7

mm long, erect. Capsules and seeds unknown. Flower-

ing time: late Oct.-mid-Nov. Figure 4.

Distribution and habitat, known only from the

Kamiesberg in central Namaqualand (Figure 3), Moraea

helmei has been collected just twice, from a “somewhat

marshy site the top of the Langkloof south of the Farm

Karas” and “from a seasonal seep south of Karas at the

upper end of the Langkloof’. Plants occur at an altitude

of about 1 125 m. Conservation status is impossible to

assess from the limited data available about the species,

but the species is undoubtedly rare.

Diagnosis and relationships', the filaments, united for

1.5 mm and slightly more than one third their length,

and the short style, ± 2 mm long, are consistent with the

M tripetala complex but the inner tepals of M helmei

are short and expanded distally into three short, subequal

lobes (vs. linear and usually cusp-like in M tripetala).

The puberulous filament column appears to be unique

in the complex. The trilobed inner tepals of M helmei

recall those of M. unguiculata, which has a longer, nar-

row, coiled central lobe and rounded lateral lobes, but

the filaments of this species are united in a prominent

column for more than half their length. A peculiarity of

the outer tepal claws of M. helmei is the pair of shallow

tooth-like projections just below the apices, a feature not

known elsewhere in Moraea'. the tepal claws in all other

FIGURE 4. — Moraea helmei, Goldblatt & Porter 13688. A, flowering

stem and corm; B, inner tepal; C, inner perianth whorl plus fila-

ments and style. Scale bar: A. 10 mm; B, C, 2 mm. Artist: John

Manning.

Bothalia42,2 (2012)

119

species are straight or slightly bowed outward and have

plane margins.

Additional specimen

NORTHERN CAPE.— 3018 (Kamiesberg): Kamiesberg, ± 1-2 km

south of Farm Karas [Welkom], in seep area west of Langkloof road,

1 125 m, (-AC), 28 Oct. 2009, Helme 3269 (NBG, photo).

3. Moraea mutila (C.H.Bergius ex Eckl.) Gold-

blatt & J.C. Manning, comb. nov. Vieusseuxia mutila

C.H.Bergius ex Eckl..- 12 (1827). Type; [Western Cape],

vicinity of Camps Bay, Bergius 15 S7238 (S, lecto.!,

designated by Goldblatt: 715 (1976b) [Syntype: [West-

ern Cape], ‘Lowenriicken’ [Lion’s Rump], Ecklon s.n.

(S, syn.!, two collections made on different dates, also

probably M mutila)].

Iris tripetala Thunb.: 13 (1782, Dec.), horn, illegit.

et nom superfl. pro M tripetala L.f. (1782, Apr.). Type:

South Africa, without precise locality, Thun berg s.n.

(UPS; Herb. Thunberg, syn.).

M tripetala van mutila Baker: 23 (1896), as nom.

nov. pro Vieusseuxia mutila Licht. ex Klatt: 621 (1866),

horn, illegit. non Eckl. (1827). Type: [Western Cape],

vicinity of Camps Bay, Bergius 15 S7238 (S, holo.!).

M punctata Baker: 1003 (1904). Type: [West-

ern Cape], ‘Piketberg Road’ [Gouda], 17 Aug. 1897,

Schlechter 4851 (K, lecto.!, designated by Goldblatt:

751 (1976); Bi, GRAI, PREI, isolecto.).

M monophylla Baker; 24 (1906). Type: [Western

Cape], ‘Olifants River, Clanwilliam’, Aug. 1894, Penther

685 [K, lecto.!, designated by Goldblatt; 751 (1976)].

Plants 15-30 cm high. Corm mostly 10-15 mm

diam., evidently without cormlets at base, tunics of

medium to coarse fibres, vertical members often thick-

ened below into prominent claws. Stem simple, rarely

1-branched (1 of 28 plants seen), glabrous or occasion-

ally pilose, sheathed below by brown, fibrous cataphylls,

these often accumulating. Foliage leaf solitary, linear,

shallowly channeled below, often becoming flat and

slightly twisted in distal third, ± half to as long as stem,

3-5(-7) mm wide, glabrous or sometimes pilose abaxi-

ally or on margins; sheathing cauline leaves 20^0 mm

long. Rhipidia mostly 2- or 3 -flowered; spathes green,

inner 35^0(^5) mm long, outer ± half as long as inner,

rarely pilose. Flowers pale blue or white, nectar guides

triangular, white to pale yellow with dark blue dots and

often edged with darker color, velvety (or smooth?);

outer tepal claws 9-12 mm long, limbs dipping at 45°,

ovate, widest in distal third, 14-15 x ± 10 mm, margins

recurving later, inner tepals ± linear to slightly expanded

in middle, sometimes as rounded lobes, 7.5-12.5 mm

long. Stamens with filaments 3. 0-3. 5 mm long, united

for 0.5-1. 5 mm; anthers 4.5-6.5(-7.5) mm long, pollen

red or yellow. Ovary narrowly ellipsoid, 7-8 mm long;

style 1-2 mm long, style branches 7-9 mm long, crests

linear, 8-10 mm long. Capsules ellipsoid, 15-17 x ± 5

mm. Seeds ± 1.8 x 1.3-1. 4 mm, pale straw-coloured,

testa spongy, angles narrowed into wings. Chromosome

number 2n= \2 (Tablet). Flowering time: late Aug.-late

Sept., depending on subspecies. Figure 5.

Distribution: largely a species of renosterveld,

Moraea mutila is centred in the western lowlands of

Western Cape, extending from the Cape Peninsula to

Piketberg, and extending locally inland to the Tulbagh

Valley (Figure 3). A collection said to be from the Olif-

ants River Valley near Clanwilliam made by Arnold Pen-

ther is more likely mislabelled and was perhaps gathered

en route from Porterville to Clanwilliam. At the very

least the Olifants River Valley record requires confirma-

tion. Soils on which M mutila has been collected are

described as shale and clay, consistent with renosterveld

but probably also on granites, and plants we have seen

grew on loamy clay. M. mutila is evidently rare on the

Cape Peninsula and as far as we have been able to deter-

mine was first collected there by C.H. Bergius in 1817,

then by Ecklon and by Zeyher in the 1 820s, and later by

Friedrich Wilms, Rudolf Marloth, and Harry Bolus but

there are no recent records. Nevertheless, it probably

persists on clay soils, its preferred habitat, on the lower

slopes of Lions Head and Signal Hill above Cape Town.

Possibly belonging here are pale blue-flowered

plants from the western lower slopes of Sir Lowry’s

Pass {Loubser 872 NBG, Goldblatt 2506 MO) with the

inner tepals trilobed as in M. mutila but the filaments are

united for ± 2.5 mm and the style is 4 mm long. Alterna-

tively these plants may be hybrids between M. tripetala

and M. unguiculata.

Diagnosis: early blooming Moraea mutila (the name

means mutilated, presumably due to the reduced inner

tepals) is recognized by the pale blue, rarely white flow-

ers with linear to narrowly lanceolate inner tepals 7.5-

12.5 mm long, erect below and spreading distally. Often

difficult to see in preserved specimens, the inner tepals

are usually slightly wider in the midline and some-

times expanded into rounded lobes near the base of the

spreading distal half. The flowers otherwise seem little

different from those of M. tripetala but the pilose nec-

tar guides are usually white or palest yellow and dotted

with dark blue to purple, whereas the nectar guides of

M. tripetala are more often yellow, edged with dark blue

FIGURE 5. — Moraea mutila. Goldblatt 2310. A, flower; B, capsule; C,

seeds. Scale bar: A, B, 10 mm; C, 2 mm. Artist: John Manning.

120

Bothalia 42,2 (2012)

or purple. The leaf is typically relatively short, in most

collections shorter than the stem, and widely channelled,

sometimes plane in the distal half. Several specimens

of two collections {Acocks 1912 and Nordenstam &

Limdgren 1998) have leaves velvety on the abaxial sur-

face and velvety spathes. The few collections from the

Cape Peninsula also have partly hairy leaves, either on

the abaxial surface or on the margins. The pubescence is

evidently variable in the populations that show the char-

acter, thus of limited value for identification, although

we have not seen any collections of true M tripetala

with pubescence of any kind on the vegetative parts.

Taxonomic history, the earliest record of Moraea

mutila is the plant described in detail by C.P. Thunberg

(Dec. 1782) under the name Iris tripetala. Thunberg’s

plant had smooth, channelled leaves, a blue flower with

yellowish nectar guides, and notably, inner tepals with

very narrow claws, convex abaxially, and patent limbs

at the base of which were two opposed lobes {denti-

bus duobus). Thunberg’s M. tripetala has, however, no

nomenclatural status or is at best superfluous as he cited

1. tripetala L.f. (April 1782) in his account. The latter,

described incompletely but said to have subulate (awl-

shaped or needle-like) inner tepals, is the basionym for

M tripetala. Thunberg actually collected multiple speci-

mens of the M tripetala complex from at least three

sites but clearly had M mutila in mind in his extended

description of /. tripetala. Since his description conflicts

with that of Linnaeus fil. we treat Thunberg’s 1. tripetala

as a separate, but superfluous name.

Moraea mutila appears again as Vieusseuxia mutila

C.H.Bergius ex Eckl. (Ecklon 1827), published with the

brief diagnosis, 'Blumen blassblau (flowers pale blue),

with citations of a C.H.Bergius collection from Camp’s

Bay (August 1817), and two Ecklon collections from

Lion’s Head [as Lowenriicken], Cape Town, one col-

lected in August (year not stated), and the other in Sep-

tember 1826. Nordenstam (1972) accepted the name as

valid despite the brief diagnosis that mentions neither

the leaf vestiture nor the form of the inner tepals, fea-

tures critical to identification of the species. The epithet

'mutila' was taken from the manuscript name on Ber-

gius’s collection, now in the Stockholm Herbarium and

annotated 'Iris nov. species mutila mihi’ in Bergius’s

hand. We wonder whether Ecklon actually saw the Ber-

gius specimen, which was probably already in Berlin

when he arrived in Cape Town in 1 823 but, nevertheless,

cited it. Bergius, incidentally, almost certainly took the

epithet 'mutila' from Martin Lichtenstein’s manuscript

Spicilegium flora capensis, which he would have seen

when he was in Berlin under Lichtenstein’s patronage at

Berlin before he departed for the Cape (Gunn & Codd

1981). The Bergius collection was designated the lecto-

type of V. mutila (Goldblatt 1976b), thus maintaining the

link between Bergius’s epithet and specimen.

Klatt’s Vieusseuxia mutila, a homonym for Ecklon’s

V. mutila, explicitly excludes the specimens on a sheet

then at the Berlin Herbarium (now at Stockholm), col-

lected by C.H. Bergius that has smooth vegetative parts

(referring them to Moraea tripetaloides DC. = M. tripe-

tala). Other specimens on that sheet with hairy leaf mar-

gins, a pilose stem and linear inner tepals are the type

of the name and are M. mutila. Baker (1896) in Flora

capensis recognized one variety of his M. tripetala, var.

mutila, with ‘leaf pilose’ [the entire diagnosis] and ref-

erences to both Vieusseuxia mutila Klatt and Iris mutila

Licht. ex Roem. & Schult. The latter, described as gla-

brous on all parts according to the protologue (Roemer

& Schultes 1817) (there is no extant type material), is in

conflict with Baker’s variety. Both name and description

of Iris mutila were taken directly from Martin Lichten-

stein’s manuscript, Spieilegium flora capensis. Klatt’s

species, however, matches Baker’s var. mutila, but as it

is illegitimate, we treat Baker’s variety as an inadvertent

new name but based on the same type as V. mutila Klatt.

Baker’s variety is M. mutila, and identical with a sec-

ond species, M. punctata, which he described in 1904,

from smooth-leaved specimens collected by Rudolph

Schlechter. Baker (1906) clearly did not realize that

his M. monophylla was a close match for M. punctata,

which now falls into synonymy under M. mutila.

Representative specimens

WESTERN CAPE. — 3218 (Clanwilliam): flats N of Piketberg at

Eendekuil turnoff, renosterveld, (-DB), 1 Sept. 1981, Goldblatt 6127

(K, MO, NBG, PRE); Piketberg, Jobskloof, (-DB), 11 Aug. 1973,

Linder 635 (BOL). 3318 (Cape Town): Dassenberg N of Mamre, W

slopes, (-AD), 23 Sept. 1974, Nordenstam & Lundgren 1998 (MO, S);

near Moorreesburg, Klein Swartfontein, renosterveld on shale, (-BA),

25 Aug. 1970, Acocks 24320 (K, MO, PRE); Porterville, flats, (-BB),

10 August 1959, Loubser 933 (NBG); slopes of Lions Rump near Tam-

boerskloof, (-CD), Aug., Zeyher 5010 (SAM); Rosebank, Cape Town,

(-CD), Aug. 1877, H. Bolus 3732 (BOL); Signal Hill, 100 m, (-CD),

Aug. 1894, Marloth 195 (PRE), 26 Aug. 1883, Wilms 3670 (K); 29

miles [45 km] from Cape Town to Malmesbury, (-DA), 29 Aug. 1932,

Lewis s.n. (BOL45082); between Salt River and Kalabas Kraal, (-DA),

4 Sept., Hutchinson 161a (K, PRE); damp flats N of Bottelary road,

(-DC), 26 Aug. 1933, Acocks 1912 (S); Stellenbosch, (-DD), 1865,

Sanderson 959 (K). 3319 (Worcester): flats S of Tulbagh Road Sta-

tion, (-AC), 11 Aug. 1974, Goldblatt 2310 (MO); Voelvlei Tortoise

Reserve, (-AC), 10 Aug. 1989, Solomon 16 (NBG); Wolseley, Farm

Romansrivier, burned April 2011, (-AC), 1 1 Aug. 2011, Boucher 7670

(MO, NBG); near Wellington, (-CC), 15 August 1926, Grant 2361

(MO).

4. Moraea cuspidata Goldblatt & J.C. Manning, sp.

nov.

TYPE. — Western Cape, 3220 (Sutherland): near Farm

Fortuin, north of Matjiesfontein, near shaded rest stop

(co-blooming with M. amabilis), (-DC), 28 Sept. 2009,

Goldblatt & Porter 13462 (NBG, holo.; MO, iso.).

Plants solitary, 180-300 mm high. Corm 15-25

mm diam., without cormlets at base, tunics of brown

to almost black, tough, medium to coarse fibres. Stem

smooth, simple or l(2)-branched exceptionally with up

to 6 branches, with cataphylls of medium textured, light

brown, mostly vertical fibres forming a collar around

base, sometimes accumulating in a dense mass. Foliage

/e<^ solitary, linear(-linear-filiform), channelled, 1. 5-3.0

mm wide, glabrous, V-shaped in section, leaf halves

closely appressed in dry conditions; sheathing cauline

leaves (30-)40-55 mm long, with dry attenuate tips.

Rhipidia several-flowered; spathes green with attenuate,

dry tips, inner (40-)50-80 mm long, outer ± half as long

as inner, glabrous. Flowers pale blue, mauve or vio-

let blue, outer tepal limbs lanceolate, with large white,

velvety nectar guides dotted and usually edged with

dark violet; outer tepal claws (8-) 10-1 2 mm, usually

speckled with dark blue to violet dots or purple in mid-

line, limbs (1 2-) 16-22 x 10-16 mm, inner tepals linear.

Bothalia 42,2 (2012)

121

(8-) 10-1 5 mm long, ascending below, distal third to half

usually spreading horizontally (or ± erect when short),

often pilose at base. Stamens with filaments {3-)4—6 mm

long, united basally for 0.5-1. 8 mm; anthers 4-8 mm

long, pollen red (rarely white). Ovary exserted, 11-15

mm long; style vestigial, branches 8-10 mm long; crests

5-8 mm long, linear, arching inward. Capsules ovoid-

ellipsoid, 14—16 mm long. Seeds 1. 8-2.0 x 1.3-1. 4 mm,

pale straw-coloured, testa spongy, thicker on angles.

Chromosome number 2n= \2 (Table 1) Flowering time:

mid-Sept. and Oct. Figure 6.

Distribution: a species of semi-arid habitats, Moraea

cuspidata grows in both mountain renosterveld and dry.

marginal fynbos, usually in sandy or sandy loam soils

derived from sandstones of the Cape System or from the

Beaufort Series of the Karoo System. It flowers late in

the season — the earliest record is in mid-September but

mid- to late October is usual. It is relatively poorly col-

lected, not (we suspect) because it is rare, but because it

grows in areas not much botanized at that time of year.

In the late spring of 2009 we found the species at several

sites we visited between Touw’s River and Komsberg

Pass in the Klein Roggeveld. The recorded range (Figure

3) extends from the Bonteberg and Voetpadsberg near

Touw’s River through the Klein Roggeveld and southern

Roggeveld to the Swartberg and higher mountains of the

FIGURE 6. — Moraea cuspidata, Goldblatt & Porter 13462. A, flowering stems and corm; B, inner perianth whorl plus filaments and style. Scale bar:

A, 10 mm; B, 2 mm. Artist: John Manning.

122

Bothalia 42,2 (2012)

Little Karoo. An outlying record from Perdekloof, south-

east of Oudtshoom near Camfer (Goldblatt & Porter

12575), probably belongs here, although the flowers are

exceptionally small, the inner tepals are 8-9 mm long,

consistent with M cuspidata. The apparent gap in the

range between the Roggeveld and the Swartberg may be

an artifact due to incomplete collecting.

Diagnosis: recognized immediately by the linear

inner tepals up to 15 mm long (rarely less than 10 mm),

spreading in the distal half (Figure 6B), Moraea cuspi-

data (named for the long, cusp-like inner tepals) also

usually has a particularly narrow foliage leaf, the leaf

halves tightly appressed in dry conditions, and brown,

sometimes almost black corm tunics, the vertical fibres

of which are usually heavily thickened. In addition,

the cataphylls tend to persist as a collar of stiff fibres

around the base (not always present in herbarium mate-

rial). The flowers are pale blue to violet with the large,

velvety nectar guides spotted with dark blue on a white

background. The filaments are usually united for 1.0-1. 5

mm but occasionally less, and ± as long as or slightly

shorter than the dark purple anthers that bear red pollen.

We have seen only two collections with ripe capsules

and these are relatively large, 14-16 mm long. The seeds

of these two collections are also relatively large for the

complex, 1. 8-2.0 mm long and unique in having a pale,

spongy testa much thickened into prominent ridges on

the angles.

The species is sympatric at some sites and even co-

blooming with the smaller-flowered Moraea amabilis

(also of the M tripetala complex), but this species has a

broadly channeled foliage leaf, sometimes plane distally

and often pilose on the abaxial surface, and consistently

short, hair-like inner tepals typically 1. 5-2.0 mm long.

At sites where we have seen the two growing together,

anthers of M amabilis have white pollen, contrasting

with the orange-red pollen of M cuspidata. The range

of M. cuspidata also coincides in part with M tripetala

subsp. violacea, which extends from Gydo Pass to the

Hex River Valley and Touw’s River, but we have not

seen them growing near one another.

Representative specimens

NORTHERN CAPE.— 3220 (Sutherland): Roggeveld, Soekop,

Huis Kamp, (-AA), 16 Sept. 2006, Rosch 556 (NBG); Koedoes Pass,

W slopes, (-CC), 22 Sept. 1970, Goldblatt 547 (BOL); Klein Rog-

geveld, Smousrand, (-DC), Sept. 2009, Goldblatt & Porter 13459

(MO, NBG); Klein Roggeveld, slopes near Farm Meintjiesplaas, (-

DC), 28 Sept. 2009, Goldblatt & Porter 13456A (MO, NBG).

WESTERN CAPE. — 3319 (Worcester): Bonteberg, Farm Kar-

rona, (-BD), 15 Sept. 1971, Thompson 1257 (NBG); 3320 (Montagu):

Bonteberg, flats at N entrance to Pienaarspoort, (-AA), 27 Sept. 2009,

Goldblatt & Porter 13453 (MO, NBG, PRE); Cabidu Karoo, Laings-

burg, (-AA), 28 Sept. 1951, Barker 7503 (NBG); Tweedside, (-

AB), ex Hort. Kirstenbosch, Barker 7466 (NBG), Sept. 1932, Lewis

s.n. Nat. Bot. Gard 2708/32 (BOL); Matjiesgoed Mtns, Farm Klein

Spreeufontein, ± 990 m, (-BC), 16 Sept. 1987, Bean, Vlok & Viviers

1938 (MO, PRE). 3322 (Oudtshoom): Swartberg Pass, rocky slopes

near picnic site, (-AC), 23 Oct. 1986, Goldblatt 8003 (MO, PRE);

top of Swartberg Pass, (-AC), Nov. 1896, H. Bolus s.n. (BOL); Cango

Caves, Oudtshoom, (-AC), 16 Oct. 1928, Gilletl 1695 (NBG); Perde-

poort N of Camfer, sandstone slopes, (-CD), 28 Sept. 2004, Goldblatt

& Porter 12575 (MO).

5. Moraea tripetala (L.f.) Ker Gawl. in Curtis’s

Botanical Magazine 19: t. 702 (1803). Iris tripetala L.f:

92 (1782).

Vieusseuxia tripetaloides DC.: 138 (1803), nom.

illegit. superfl. pro I. tripetala L.f. (1782). Vieusseuxia

tripetala (L.f) Voigt: 602 (1845). Vieusseuxia tripetala

(L.f) Klatt: 155 (1894), horn, illegit., non (L.f) Voigt

(1845). Type: [Western Cape], without precise locality,

Thunberg s.n. Herb. Thunb. 1186 [UPS-THUNB, lecto.!

designated by Goldblatt: 175 (1976)].

Plants (140-)250^50 mm high, glabrous on all

vegetative parts. Corm mostly 8-15 mm diam., with-

out cormlets at base, tunics of moderately coarse,

wiry, usually dark grey fibres. Stem simple or 1- or

2(-6)-branched, sheathed below by brown cataphylls

and sometimes with collar of brown fibres. Foliage

leaf solitary (rarely 2 in subsp. violacea), leathery, nar-

rowly channeled, C- or V-shaped in section, to 5 mm

wide, usually exceeding stem and distally trailing;

sheathing cauline leaves mostly 45-60 mm long. Rhip-

idia 3-several-flowered; spathes green with dry attenu-

ate tips, inner 35-65(-70) mm long, outer ± half as long

as inner. Flowers pale blue, purple, or violet, nectar

guides triangular, yellow edged with violet, or white dot-

ted with dark blue and edged with darker blue, lightly

honey scented, unscented or strongly scented of carna-

tion (subsp. violacea)-, outer tepals 20-30(-32) mm

long, claws 9-12 mm long, limbs obovate, widest in dis-

tal third, 11-18 x ± 8-13 mm, plane, spreading at 45°,

margins eventually recurved, inner tepals hair-like to ±

linear, l-4(-6) mm long, or lacking. Stamens with fila-

ments 4-6 mm long, united for 0.5-1. 5 mm, occasion-

ally (southern populations) ± free; anthers 4-7 long,

pollen usually orange-red, rarely white. Ovary 4-9 mm

long; style 1-2 mm long, style branches 9-12 mm long,

crests ± linear, 5-12 mm long. Capsules ellipsoid, 8-14

mm long. Seeds 1.0-1. 3 mm x ± 1 mm, flat surfaces

slightly wrinkled, brown seed body visible through testa,

angles forming raised, golden-brown ridges. Chromo-

some number 2n = \2 (only subspp. tripetala and vio-

lacea counted) (Table 1). Flowering time: mainly Aug.-

mid-Sept., occasionally in Oct. Figures 7, 8.

Distribution: widespread in Western Cape, Moraea

tripetala extends from Aurora on the west coast to near

Knysna in the east, and inland to Ceres and the Little

Karoo (Figure 9). Habitats vary from limestone flats in

fynbos, to neutral and acid sands in sandveld, to loamy

alluvium in coastal and montane fynbos. Interior popu-

lations from the Warm Bokkeveld (Ceres) and western

Little Karoo, here segregated as subsp. violacea, occur

in significantly drier habitats on clay soils in renos-

terveld. Populations segregated as subsp. jacquiniana

are mostly montane and occur above 500 m but at lower

elevations in the southern Cape Peninsula, and flower

significantly later in the year. Although much of its low-

land habitat has been lost to agriculture and its original

range significantly reduced, M. tripetala still occurs

extensively in undisturbed low to middle elevation sites.

Interior populations have lost little of their original habi-

tat except immediately around Ceres where orchards

have replaced much of the original vegetation.

Diagnosis: in our revised, narrower circumscription,

Moraea tripetala includes plants with reduced hair-like

to linear inner tepals mostly 2-A, rarely up to 6 mm

long, and filaments united for up to 1.5 mm and occa-

sionally free. The outer tepal limbs are broadly ovate.

Bothalia 42,2 (2012)

123

FIGURE l.—Moraea tripetala subsp. tripetala. A, B, D, Goldblatt & Porter 13467 (MO); C, Goldblatt & Porter 12374. A, flowering stems and

corm; B, C, inner perianth whorl plus filaments and style; D, seeds. Scale bar: A, 10 mm; B-D, 2 mm. Artist: John Manning.

124

Bothalia 42,2 (2012)

FIGURE 8. — Moraea tripetala suhsp. jacquiniana. Red Hill, no voucher.

Scale bar; 10 mm. Artist: John Manning.

longer than wide and oriented ± 45° below horizontal.

The corms have dark brown to grey tunics, usually com-

posed of medium-textured fibres, and do not produce

cormlets at the base. The linear leaves, normally exceed-

ing the stem, are narrowly channeled, and in dry condi-

tions the leaf halves lie parallel to one another. As far as

we can determine, vegetative reproduction by produc-

tion of cormlets does not occur in M tripetala as now

circumscribed. Flower colour ranges from pale to deep

blue or violet or rarely light purple, sometimes even in

the same population. Nectar guides are often yellow

edged with dark blue or purple, but occasionally white,

then dotted with blue to purple. The seeds are small, and

typical of the complex in being 5(6)-sided with the fac-

ets slightly wrinkled and showing the brown color of the

seed body (Figure 7D). The angles are raised into nar-

row wings of pale straw colour. In contrast, M amabilis,

M. cuspidate, and M grandis have larger seeds of more

complex structure.

Variation: there are a number of notable variants

among the plants we include in Moraea tripetala, one

of the most important of which includes populations

from the interior Western Cape on clays and shales from

Ceres to the western Little Karoo. These plants have

dark violet outer tepals with yellow nectar guides, are

usually relatively short in stature, and have short stiff,

hair-like inner tepals, mostly ± 2 mm long. The flow-

ers we have examined alive have a strong clove or car-

nation-like scent. Unusually for the complex, and for

subg. Vieusseuxia as a whole, some populations of this

morph have two foliage leaves (e.g. Dymond s.n. from

near Ouberg Pass; Goldblatt 4184, 11438 from Op-de-

Tradouw near Barrydale; Van Wyk 61 from the Voetpads-

berg; Goldblatt & Snijman 697 6 A from Worcester). We

recognize these populations of the interior Western Cape

as subsp. violacea.

Elsewhere, Moraea tripetala has pale or deep blue,

or sometimes purple outer tepals, either with yellow or

white nectar guides, in the latter case usually speckled

with dark blue or purple, and flowers are faintly sweet-

scented or odourless. It is also relatively tall (allowing

for reduced stature in years of poor rainfall) and usu-

ally has hair-like to linear inner tepals 2-4(-6) mm long

(occasionally the inner tepals are absent in southern

Cape populations). Among these populations, those that

flower from November to January on the Cape Penin-

sula and surrounding mountains stand out in their softly

fibrous corm tunics, slightly smaller flowers, and usually

very narrow leaves. The flowers also often have inner

tepals 4-6 mm long. Typical M. tripetala in the same

area flowers from August to early October. We think it

useful to recognize these two sets of populations taxo-

nomically but favour subspecies rank because the mor-

phological differences between them and typical M. tri-

petala are small and not absolute. The taxon was treated

by Lewis (1941, 1950) as war. jacquiniana.

Variation elsewhere across the wide range of Moraea

tripetala seems to us less taxonomically significant and

less consistent. We discuss this under subsp. tripetala.

History: although the type of Moraea tripetala was

collected by C.P. Thunberg, the basionym Iris tripetala

was described by Linnaeus fil. (April, 1782), several

months before Thunberg (Dec. 1782) published his own,

much more extensive description. Goldblatt (1976b)

selected as lectotype a specimen that had short, hair-like

inner tepals and a long, narrow, channelled leaf, thus

fixing the application of the name. Thunberg’s plant is

obviously a different species, and has the distinctive

inner tepals of what is now M mutila (see extended dis-

cussion under that species). The reduced inner tepals

rendered M. tripetala so distinctive that it acquired few

synonyms. De Candolle (1804), when transferring /.

tripetala to Vieusseuxia, the genus then used for south-

ern African species originally treated as Iris, renamed it

V. tripetaloides, a morphologically more apt epithet. The

name is superfluous as De Candolle cited 7. tripetala L.f.

Bothalia 42,2 (2012)

125

16° 18° 20° 22° 24° 26° 28°

FIGURE 9. — Distribution of Moraea tripetala subsp. tripetala, •; M.

thpetala subsp. violacea, A; M tripetala jacqidniana, o.

and I. tripetala Thunb. in synonymy. I. tripetala was

transferred to Moraea in 1803 by Ker Gawler but the

confusion over the circumscription of Moraea persisted

and Voigt (1847) and then Klatt (1866) independently

transferred /. tripetala to Vieusseuxia.

Key to subspecies

la Plants mostly of montane habitats in rocky sandstone soils,

flowering mainly November to January, occasionally in

October; ovary 4—6 mm long; inner tepals often 5-6 mm

long, reaching bases of anthers; tepals mostly dark violet

5c. suhs'p. jacquiniana

lb Plants of clay, granitic or sometimes rocky limestone or

sandstone soils, flowering mainly August to early Octo-

ber; ovary 6-9 mm long; inner tepals l-3(-5) mm long (or

absent), rarely reaching base of anthers; tepals pale to dark

blue to violet:

2a Ovary 6-8 mm long; outer tepals 24—26 mm long and

inner tepals 1-2 mm long; tepals violet with yellow nec-

tar guides; foliage leaves 1 or 2; plants of clay soils in

renosterveld in the Warm Bokkeveld, Bonteberg, and

Little Karoo 5b. subsp. violacea

2b Ovary 8-9 mm long; outer tepals 23-30(-32) mm long;

and inner tepals absent or l-3(-5) mm long; tepals pale

blue or deep blue or purple usually with white (occasion-

ally yellow) nectar guides; foliage leaf always solitary;

plants of sandy flats and slopes, sometimes in limestone,

extending from Aurora to Knysna 5a. subsp. tripetala

5a. subsp. tripetala

llris mutila Licht. ex Roem. & Schult.: 447 (1817).

Vieusseuxia mutila (Licht. ex Roem. & Schult.) A.Dietr.:

494 (1833), horn, illeg. non Eckl. (1827). Type;

unknown, assigned here based on description alone.

Plants 250-450 mm high, stem sometimes sheathed

below by collar of brown fibres. Corm mostly 12-15

mm diam. Foliage leaf always solitary, linear. Inner

spathes 45-65(-70) mm long. Flowers pale blue or pur-

ple with yellow or white nectar guides, faintly honey-

scented or unscented; outer tepals 23-30(-32) mm long,

claws 10-12 mm long, limbs 12-18 mm long,; inner

tepals usually hair-like, 1-3 (-5) mm long, or absent.

Stamen filaments 4-6 mm long, united for 1.0-1. 5 mm,

occasionally (southern populations) ± free; anthers

4. 5-7.0 long; pollen usually red or white. Ovary 8-9

mm long; style branches ±12 mm long, crests mostly

8-12 mm. Capsules 12-14 mm long. Flowering time:

late Aug.-mid-Sept., occasionally in Oct. (exceptionally

July, e.g. Wurts 212). Figure 7.

Distribution: largely a plant of coastal forelands,

subsp. tripetala extends from Aurora on the west coast

of Western Cape to near Knysna in the east. It also

occurs on lower mountain slopes, occasionally up

to ± 700 m, and has been recorded on the Piketberg,

the Hottentots Holland-Jonkershoek mountain com-

plex, the Witzenberg at Ceres, and the lower southern

slopes of the Langeberg (Figure 9). Populations along

the southern Cape, notably in the De Hoop area, the

Agulhas Peninsula {Goldblatt 2607), and some from

near Caledon {Goldblatt 6196) occasionally lack inner

tepals, or individual flowers may lack one or two of

the inner tepals (e.g. Goldblatt & Manning 12248, east

of Swellendam). Plants from the southern foothills and

lower slopes of the Langeberg sometimes have free fila-

ments (e.g. Thorne s.n. SAM, from Garcia’s Pass; Gold-

blatt 3741 from near Suurbraak). Subsp. tripetala is usu-

ally found in sandy habitats or on coastal limestone, thus

in sandveld, strandveld and fynbos vegetation.

Representative specimens

WESTERN CAPE. — 3218 (Clanwilliam); Piketberg, plateau

on Kapteins Kloof Mtn, (-DA), 21 Oct. 1935, Pillans 7796 (BOL);

Piketberg, plateau E of Levant Mt., edge of dam, (-DD), 20 Sep-

tember 1974, Linder 414 (BOL). 3318 (Cape Town): R27 to Lange-

baanweg, sandveld, (-AC), 29 Sept. 2009, Goldblatt & Porter 13467

(MO); Dassenberg, N of Mamre, (-AD), 23 Sept. 1974, Nordenstam

& Limdgren 1999 (MO, S); near Mamre Road Station, sandy water-

logged ground, (-BC), 3 Sept. 1974, Goldblatt 2487 (MO); sandy

flats near Wynberg, (-CD), Sept. 1884, Macowan 215 (BOL); Camps

Bay, (-CD), Sept. 1886, Thode 8531 (NBG). 3319 (Worcester): Ceres

Nature Reserve, top of MichelTs Pass, rocky sandstone slopes, (-CD),

29 Sept. 2009, Goldblatt & Porter 13463 (MO, NBG). 3320 (Mon-

tagu): Swellendam, below Langeberg range at Clock Peaks, (-CD), 2

July 1952, Wurts 212 (NBG). 3419 (Caledon): Caledon Swartberg and

the Baths, (-AB), Aug., Ecklon <& Zeyher s.n. (MO); 18.4 km SW of

Greyton, shale ground in renosterveld, (-BA), 28 August 1970, Acocks

24337 (K, MO, PRE); Rotary Drive, Hermanns, (-AC), 10 October

1974, Goldblatt 3009 (MO, PRE); hill above. Pearly Beach, (-CB), 11

Sept. 1974, Goldblatt 2607 (K, MO, PRE). 3420 (Bredasdorp): Suur-

braak to Heidelberg, W of Strawberry Hill, (-BA), Goldblatt 3741

(MO); clay bank E of Swellendam, (-BB), 7 Sept. 2003, Goldblatt &

Manning 12248 (MO, NBG); 10 km S of Bredasdorp, entrance to Die

Poort, limestone, (-CA), 13 Sept. 1978, Goldblatt 4860 (MO). 3421

(Riversdale): Riversdale, Garcias Pass, (-AA), Oct. 1926, in Thorne

s.n. (SAM 43214); 2 km from Herbertsdale to Cloete’s Pass, (-BA),

26 Sept. 2003, Goldblatt & Porter 12374 (MO). 3422 (Mossel Bay):

Pacaltsdorp, George, (-AB), 22 Aug. 1978, Moriarty 339 (MO); Gou-

kamma Nature Reserve, (-BB), 1968, Heinecken 206 (PRE).

5b. subsp. violacea Goldblatt c& J.C. Manning, subsp.

nov.

TYPE. — Western Cape, 3320 (Montagu): Op-de-

Tradouw, Barrydale to Montagu, south facing clay slope

in renosterveld, (-DC), 26 Aug. 2000, Goldblatt 11438

(NBG, holo.; K, MO, PRE, iso.).

Plants mostly 200-300 mm high, stem sometimes

with collar of fine fibres around base. Corm mostly

8-10 mm diam., tunics of medium-textured, usually

pale fibres, sometimes thickened below. Foliage leaf 1

or 2, linear, mostly 1.5-2. 5 mm wide, leaf halves often

folded together and blade appearing terete. Inner spathes

35^5(-50) mm long. Flowers violet with yellow, vel-

vety nectar guides, strongly seented of cloves (?always);

outer tepals 24-26 mm long, claws ± 10 mm long, limbs

14—16 mm long, inner tepals hair-like, 1-2(3) mm long.

Stamen filaments 4. 0-6. 5 mm long, united for 0.5-1. 2

mm; anthers 4. 0-6. 5 mm long, pale violet, pollen white.

126

Bothalia 42,2 (2012)

Ovary 6-8 mm long; style branches ± 12 mm long,

crests 6-8(10) mm long. Capsules 8-11 mm long. Flow-

ering time: late Aug.-late Oct.

Distribution', subsp. violacea extends from Gydo Pass

and the Warm Bokkeveld though the southern foothills

of the Hex River Mtns near Worcester inland to the

Bonteberg, Touws River and the western Little Karoo

near Barrydale (Figure 9). Populations with two foliage

leaves appear to occur randomly across its range. Plants

almost invariably occur on clay soils in renosterveld but

we have also seen the species growing among low suc-

culent shrubs.

Representative specimens

WESTERN CAPE.-3319 (Worcester): Gydo Pass, (-AB), Barker

6836 (NBG); Hottentots Kloof, (-BA), 28 Sept. 1944, Barker 3031

(NBG); E slopes of Theronsberg Pass, (-BC), 31 Oct. 1974, Gold-

blatt 3228 (MO); Ceres, Lakenvlei, (-BC), 10 Oct. 1941, Barker 1336

(NBG); 2 km east of De Dooms, Hex River Valley, (-BC), 27 Sept.

1974, Nordenstam <£ Lundgren 2056 (MO, S); ± 12 km N of Worcester

at Brandwagt turnoff, Worcester West, (-CB), 26 Sept. 1983 (fr.), ex

hort. Missouri, Mar. 1986, Goldblatt & Snijman 6976A (MO); Doring-

kloof, Voetpadsberg, (-DB), 24 Aug. 1985, Van Wyk 61 (NBG); Koo,

foot of Naudesberg, (-DD), 12 Sept. 1962, Lewis 6054 (NBG). 3320

(Montagu): Ouberg to Touws River, (-CB), 29 Sept. 1932, Dymond

s.n. {BOL21234); Keurkloof, off Cogman’s Kloof, (-CC), 24 Sept.

1935, Levi’is s.n. (BOL); Op-de-Tradouw, Barrydale to Montagu, S fac-

ing clay slope in renosterveld, (-DC), 23 Sept. 1976, Goldblatt 4184

(MO), 12 Sept. 1994, Goldblatt & Manning 9990 (K, MO, NBG).

5c. subsp. jacquiniana (Schltr ex G.J. Lewis) Gold-

blatt & J.C. Manning, stat. et comb. nov. Moraea tri-

petala var. jacquiniana Schltr. ex G.J. Lewis: 54 (1941).

Type: [Western Cape], Constantiaberg, Dec. [without

year], Wolley Dod 1919 (BOL, lecto.!, designated by

Goldblatt: 751(1976b); K!, iso.).

Vieusseuxia pulchra Eckl.: 13 (1827). Type: [Western

Cape], ‘Hottentotshollandkloof’, 25 Nov. [without year],

Ecklon & Zeyher s.n. (S, holo.!).

Plants 140-300 mm high, stem usually sheathed

below by collar of fine, loosely netted fibres. Corms ±

10 mm diam., tunics of relatively fine fibres, not usu-

ally accumulating. Foliage leaf solitary, narrowly chan-

nelled or leaf halves appressed together thus appearing

terete, exceeding stem and up to twice as long, 2-3(-5)

mm wide (when dry often apparently ±1.5 mm diam.).

Inner spathes 40-50 mm long. Flowers usually purple

or blue with triangular, white nectar guides spotted with

dark blue and edged darker color, unscented, outer tepals

20-25 mm long, claws 9-11 mm long, limbs lanceolate,

11-16 X 8-11 mm, inner tepals hair-like, mostly 5-6

mm long, reaching or exceeding anther bases. Stamen

filaments 4-6 mm long, united for 1. 0-1.5 mm; anthers

5-6 long, pollen usually (always?) red. Ovary 4—6 mm

long; style branches 10-11 x 2 mm long, crests linear,

5-8 mm long. Capsules mostly 10-14 mm long. Flower-

ing time'. mid-Nov.-Jan., rarely in late Oct. Figure 8.

Distribution', occurring entirely within the range of

subsp. tripetala, subsp. jacquiniana extends from the

Cape Peninsula to the Hottentots Holland and Houw

Hoek Mtns and north to the mountains around Fransch-

hoek, Paarl, and Wellington (Figure 9). There are also

isolated records from the Piketberg, the mountains near

Ceres, and in the vicinity of Citrusdal {Ecklon & Zey-

her Irid 20), this last in need of confirmation. Although

largely montane, subsp. jacquiniana occurs at quite low

elevations in the southern Cape Peninsula. Plants grow

in fynbos on stony sandstone slopes. Flowering is very

late in the season, mostly November and December, but

plants have been collected in January and sometimes

in late October. Subsp. tripetala flowers in August and

September, usually at lower elevations, sometimes in

similar habitats.

Diagnosis', snh?,^. jacquiniana is poorly differentiated

from subsp. tripetala but apart from the later flowering

time it can be distinguished by the finely fibrous corm

tunics, shorter rhipidial spathes 40-50 mm long, and

slightly smaller flowers, usually deep violet, consistently

with inner tepals 4—6 mm long (usually shorter or absent

in subsp. tripetala). The outer tepals are typically 20-25

mm long, the ovary 4—6 mm long and capsules, 10-14

mm long, all smaller than in subsp. tripetala. As far as is

recorded, the violet flowers consistently have white nec-

tar guides whereas M. tripetala has yellow or less often

white nectar guides, then speckled with blue dots. The

colour illustrations of flowers of the two taxa can read-

ily be compared in Wild Flowers of the Cape Peninsula

(Maytham Kidd 1950).

The taxon was named in honor of N.J. Jacquin, whose

monumental volumes of coloured illustrations of plants,

many of them from the Western Cape and Namaqua-

land, were an inspiration and major resource for accurate

plant depictions, including his painting of three variants

of Moraea tripetala {leones plantarum rariorum vol. 2,

t. 211), one of which perhaps was thought by Schlech-

ter to represent what is now subsp. jacquiniana. The

epithet was first used, without description, by Bolus &

Wolley Dod (1903) in a checklist of plants of the Cape

Peninsula as if validly published by Schlechter. The epi-

thet was taken from the collection labels of Schlechter

7222 (collected in 1 896) and was only validated at vari-

etal rank by G.J. Lewis in 1941. A collection from ‘Hot-

tentots Holland Kloof’ was named Vieusseuxia pulchra

by Ecklon in 1827. Ecklon’s descriptive phrase, ‘flowers

darker blue’ [than M. setacea the name Ecklon used for

V. tripetala] barely qualifies as a diagnosis but Norden-

stam (1972) regarded the name as validly published.

Representative specimens

WESTERN CAPE.— 3218 (Clanwilliam): along the Olifants

River, Clanwilliam to Citrusdal, (-?BB), Nov., Ecklon & Zeyher Irid

20 (MO); Piketberg, plateau on Kapteins Kloof Mtn, (-DA), 21 Oet.

1935, Pillans 7796 (BOL). 3318 (Cape Town): Cape Peninsula, Table

Mt., (-AD), Dec. 1950, Pillans 10279 (BOL, MO); Table Mt., lower

plateau, Nov. 1944, Lewis 946 (SAM). 3319 (Worcester): Michell’s

Peak, Ceres, (-AD), 16 Dec. 1948, Esterhuysen 14767 (BOL); Upper

Wellington Sneeukop, moist slope below shale band, flowers dark

violet-blue, (-CA), 23 Jan. 1972, Esterhuysen 32799a (BOL); Groot

Drakenstein Mtns, Devil’s Tooth, (-CC), 12 Dec. 1943, Esterhuysen

9549 (BOL); mountain at top of Franchhoek Pass, (-CC), 18 Nov.

1974, Nordenstam & Lundgren 2259 (MO, NBG); Kaaimansgat, neck

above High Noon Estate, (-CD), 6 Jan. 1980, Goldblatt 5422 (MO);

Onklaarberg, 20 miles [± 30 km] S of Woreester, (-DC), Dee. 1924,

Stokoe 1106 (PRE). 3418 (Simonstown): Cape Peninsula, Vlakken-

berg, (-AB), 1 Jan. 1896, Wolley Dod 457 (BOL); burned lower slopes

of Klaasjagersberg adjaeent to Cape Point Reserve, (-AB), 26 Nov.

1979, Goldblatt 5258 (MO, PRE); Paulsberg slopes, (-AB), 6 Nov.

1939, Lewis 675 (SAM). Muizenberg Mt., 1 500 ft [465 m], (-AB), 29

Nov. 1938, Wall s.n. (S); Sir Lowry’s Pass, 1 219 m, ' Moraea jacquini-

ana Schltr., n. sp.’, (-BB), 14 Jan. 1896, Schlechter 7222 (GRA, PRE).

3419 (Caledon): Haasvlakte, Houwhoek road to Highlands, (-AA),

Bothalia 42,2 (2012)

127

FIGURE 10. — Moraea hainebachi-

ana, Goldblatt & Porter

13262. A, flowering stems

and corm; B, inner perianth

whorl plus filaments and style.

Scale bar: 10 mm. Artist: John

Manning.

14 Oct. 1988, Boucher & Stindt 5452 (NBG, PRE); Lebanon Forest

Reserve, Jakkalsrivier Catchment la, (-AA), 23 Nov. 1970, Haynes

476 (NBG, PRE).

6. Moraea hainebachiana Goldblatt & J.C. Manning,

sp. nov.

TYPE. — Western Cape, 3217 (Vredenburg):

Jacobsbaai, 1 km E of town on calcrete ridge, Erf 890,

(-DD), 31 Aug. 2011, Claassens 95 (NBG, holo.; MO,

iso.).

Plants 180-280 mm high. Corm mostly 7-12 diam.,

usually with cormlets at base, tunics of coarse, almost

black, hard, wiry fibres, thickened below into claw-like

ridges. Stem usually simple, rarely 1 -branched, glabrous,

bearing several, dark-coloured cormlets just below

ground level in foliage leaf axil. Foliage leaf solitary,

linear, channelled, inserted well above top of cataphyll,

trailing above, (1.5-)2.0^.0 mm wide, exceeding stem

by 100-150 mm; sheathing cauline leaves ± 55 mm

long, green with dry attenuate apices, lowermost some-

128

times with axillary coimlets. Rhipidia usually 2-flow-

ered, spathes green with dry attenuate tips, inner 60-65

mm long, outer ± half as long as inner. Flowers pale vio-

let to deep blue, outer tepal limbs darkly veined, nectar

guide white with lines of dark blue radiating from limb

base, scented of vanilla, outer tepals 24—25 mm long,

claw ± 10 mm long, white-hairy on adaxial surface, with

prominent basal, yellow nectary ± 1.5 mm long, limb

14-15 X 9 mm, usually ultimately recurving, inner tepals

spindle-shaped, sometimes oblanceolate and obscurely

3-lobed, 5-6 x 0.5 mm, tapering distally with apex

curving inward, sometimes inner distal surface pilose.

Stamens with filaments ± 6 mm long, united in lower

1.0-1. 5 mm; anthers 5-6 mm long, blue-grey, pollen

off-white or yellow. Ovary narrowly oblong-truncate,

included or exserted, 7-9 mm long; style ± 2 mm long,

dividing ± 1 mm above top of filament column, branches

wedge-shaped, 7-8 x ± 1.5 mm at apex; stigma shal-

lowly bilobed; style crests narrowly wedge-shaped (or

sublinear), ±10x1 mm. Capsules not developed, shed

soon after flowers wilt. Seeds not produced. Chromo-

some number unknown. Flowering time: Aug.-mid Sept.

Figure 10.

Distribution: restricted to the Saldanha District in

Western Cape (Figure 11), Moraea hainebachiana is a

narrow edaphic endemic of rocky, limestone flats and

slopes and calcareous sands along the coast and adja-

cent hills. Plants grow in humus-rich pockets of loam

between fractured limestone as well as in coarse calcare-

ous sand. Populations extend from the Farm Trekoskraal

north of Jacobsbaai through the limestone hills north of

Saldanha to the southern end of the Donkergat Peninsula

in West Coast National Park. Distinctive as the habitat

is, typical M tripetala grows in the same places, but

blooms four to five weeks later. M hainebachiana was

evidently first collected in 1932 near Langebaan by the

late G.J. Lewis, expert on the systematics of southern

African Iridaceae. Her collection was referred with-

out question to M tripetala at the time. The probabil-

ity that the species was distinct from M tripetala was

brought to our attention by Koos (Jakobus) Claassens of

Jacobsbaai, who has made a thorough study of the local

and strongly endemic flora of the Saldanha limestone

areas. Given its very narrow range, M hainebachiana

must be considered threatened by coastal development

although it is probably secure at the southern end of its

FIGURE 1 1 . — Distribution of Moraea amahilis, •; M hainebachiana,

A; M. ogamana, o.

Bothalia 42,2 (2012)

range in the West Coast National Park under current low

wildlife stocking levels.

Diagnosis: named for the Hainebach family of Cape

Town for their generous contribution to conservation

of the Cape flora, Moraea hainebachiana at first seems

to be fairly typical M tripetala except for its low stat-

ure and slightly smaller flowers. On close examination,

however, the species exhibits several unusual features.

The first of these is the cluster of small, dark grey or blu-

ish cormlets in the foliage leaf axil and at the base of the

corm, the latter feature especially pronounced in non-

flowering individuals. The foliage leaf is always inserted

on the stem well above the top of the cataphyll and

clearly separated from it whereas in other species in the

complex, the insertion of the foliage leaf is concealed by

the cataphyll. The flowers of M hainebachiana are con-

sistently pale violet to deep blue and have inner tepals

5-6 mm long, somewhat longer than is typical for M

tripetala and unusual in being expanded in the mid-

dle, thus spindle-shaped or sometimes oblanceolate and

obscurely 3-lobed, and occasionally pilose on the inner

(adaxial) surface. More significantly, the filaments are as

long as or slightly longer than the anthers, whereas the

anthers are slightly longer than the filaments in M trip-

etala and in some species of the complex considerably

so. The anthers are pale grey-blue and contain whitish

or yellow pollen. Orange-red pollen is more frequent in

the complex. The nectar guides of M hainebachiana are

also somewhat unusual, consisting of lines of dots on a

whitish background with the edges not clearly defined

as they are in M tripetala in which the yellow or white

nectar guides are typically edged in darker blue to violet.

Vegetative reproduction does not occur in M. tripetala as

circumscribed here and the production of multiple corm-

lets in leaf axils and at the base of the corm is unknown

elsewhere in the M. tripetala complex.

Visiting the type locality at Jacobsbaai after flowering

to examine capsules and seeds, we found all capsules

poorly developed and ready to be shed, if not already

fallen, before developing any seeds. We confirmed the

condition for two more populations. Microscopic exami-

nation of pollen grains shows them to be malformed and

evidently infertile. We conclude that M. hainebachiana

is a vegetative apomict. Propagation is effected by corm-

lets that plants liberally produce.

Representative specimens

WESTERN CAPE. — 3217 (Vredenburg): Jakobsbaai, Swart-

riet Farm, calcrete ridge to N of road to farm house, 50 m, (-DD), 7

Aug. 1993, Boucher 5801 (NBG); Jacobsbaai, limestone pavement,

(-DD), 1 Sept. 2010 (ft.), Goldblatt & Manning 13491 (MO, NBG,

PRE); limestone hill N of Saldanha, limestone pavement, (-DD), 4

Sept. 2009 (fl,), Goldblatt <6 Porter 13262 (MO). 3318 (Cape Town):

Langebaan, (-AA), 4 Sept. 1932, Lewis s.n.BOL45094 (BOL); West

Coast National Park, flats SW of Konstabel Kop on calcareous sands,

(-AA), 13 Sept. 2011, Manning & Goldblatt 3338 (NBG).

7. Moraea ogamana Goldblatt & J.C. Manning, sp.

nov.

TYPE. — Western Cape, 3319 (Worcester): Elands-

berg Farm, foot of Elandskloof Mts, seasonally wet allu-

vial flats, in fynbos, (-AC), 9 Sept. 2010, Goldblatt &

Manning 13520 (NBG, holo.; MO, iso.).

Bothalia 42,2 (2012)

129

Plants 150-200(-380) mm high. Corm 6-12 mm

diam., tunics of hard, wiry, black fibres. Stem usually

unbranched, 2 or 3 intemodes long. Foliage leaf soli-

tary, sometimes with second leaf from axillary cormlet,

usually shorter than stem, rarely longer, 2-3 mm wide

when opened flat, widely channelled or leaf halves

almost folded together, smooth, apple-green, inserted at

ground level or shortly below ground, usually bearing

small axillary cormlet; sheathing cauline leaves (30-)45

mm long, green with dry attenuate tips. Rhipidia mostly

3 -flowered; spathes green with dry brown attenuate tips,

inner to 55 mm long, outer ± half as long as inner. Flow-

ers pale blue, with triangular yellow nectar guides, with

dark veins radiating from nectar guide, outer tepal claws

12-14 mm long, limbs ovate, ± 15 mm long, inner tepals

hair-like, up to 4 mm long. Stamens with filaments ±

5 mm long, united for ± 1 mm; anthers ± 6 mm long,

dark bluish black, pollen white. Ovary linear-cylindric,

10-13 mm, darkly lined vertically on locules and septa;

style ± 2 mm long, branches ±12 mm long, crests ± lin-

ear, 8-12 mm long. Capsules ± cylindric, (15-)20-24 x

2 mm. Seeds 0.8-1. 5 mm diam., golden brown, shortly

winged on angles. Chromosome number unknown.

Flowering time'. mid-Aug.-mid-Sept. Figure 12.

Distribution: restricted to the Western Cape low-

lands between Voelvlei and Strand and locally in the

upper Breede River Valley between Wolseley and Botha,

Moraea ogamana occurs in waterlogged stony ground

FIGURE 12. — Moraea ogamana, Goldblatt & Manning 13520. A, flow-

ering stem; B, capsules; C, seeds. Scale bar: 10 mm. Artist; John

Manning.

(Figure 11). Judging by the few collections, the species

is rare. With rapid agricultural development in lowland

Western Cape M ogamana must be regarded as endan-

gered, although its existence is secure in the Elandsberg

Nature Reserve at Bo-Hermon immediately south of

Voelvlei, a small part of its once much wider range.

Diagnosis: the narrow, linear-cylindric ovary 10-13

mm long, often partly included in the spathes, and pale

blue flower with outer tepal limbs with yellow nec-

tar guides and dark radiating venation immediately set

Moraea ogamana apart in the M. tripetala complex.

Associated with these features are the short stature,

rarely exceeding 180 mm, fairly small corms, 7-12 mm

diam., with wiry, black tunic fibres and a relatively short

leaf, usually shorter than, but occasionally exceeding

the stem. Unlike most populations of M. tripetala in the

Western Cape forelands, which have orange-red pollen,

the pollen of M ogamana is white. An additional, but

somewhat trivial distinction is the style crests, up to 12

mm, which are unusually long for the relatively small

flowers. More importantly, almost all specimens we

have seen have a single, small cormlet in the foliage leaf

axil. This modest level of vegetative reproduction results

in plants sometimes growing in small clumps of three or

four individuals. Correlated with the long, angular ovary,

the capsules are ± cylindric and normally 20-24 mm

long. Capsules of M tripetala are ellipsoid-oblong and

only 10-14 mm long. The species is named in honour of

Ms Naoka Ogama of Japan for her generous donation to

the study of southern Afiican Iridaceae.

Representative specimens

WESTERN CAPE.-3318 (Cape Town): between Paarl and Pont,

(-DB), Drege s.n. (S); near Groenfontein, Klapmuts, level wet area,

(-DD), 15 Sept. 1983, Van Zyl 3515 (NBG, PRE); damp place near

Bottelary Road, (-DD), 9 Sept. 1934, Acocks 2157 (S). 3319 (Worces-

ter): Bo-Hermon, Elandsberg Farm, seasonally wet alluvial fynbos, (-

AC), 20 Oct. 2010 (fr.). Manning 3307 (MO, NBG); between Bain’s

Kloof and Wolseley, wet sandy flats, (-AC), 24 Aug. 1974, Goldblatt

2428 (MO); Worcester, Botha’s Halt, (-CB), 14 Sept. 1928, Gillett

293 (NBG). 3420 (Simonstown): Harmony Reserve, Strand, (-BB), 28

Aug. 2000, Runnalls 1041 (NBG).

8. Moraea amabilis Diels in Botanishe Jahrbticher

der Systematik 44: 118 (1910). Type: [Northern Cape],

Bokkeveld, Calvinia, Oorlogskloof, 13 Sept. 1900, Diels

626 (B, lecto.!, designated by Goldblatt: 751 (1976), as

holotype).

Plants 150-250 mm high, growing in clonal colo-

nies. Corm mostly 7-12 mm diam., with tunics of pale,

medium to fairly coarse fibres (rarely soft relatively

fine fibres), often replaced annually by two daughter

corms. Stem simple or l(2)-branched, glabrous or vel-

vety, sometimes with cormlets in below-ground axils,

enclosed at base by brown cataphylls, occasionally accu-

mulating in a collar of fine fibres around base. Foliage

leaf solitary, linear, channeled, usually broadly so, 2-5

mm wide, abaxial surface often velvety-pilose; cauline

sheathing leaves ± 40-52 mm long, glabrous or vel-

vety, green with dry apices. Rhipidia several-flowered;

spathes green, dry and attenuate above, becoming dry

entirely, glabrous or velvety, inner mostly 45-55 mm

long, outer ± half to two thirds as long as iimer. Flow-

ers mostly purple or violet to dark blue (rarely pink,

dull yellow or buff-brown), nectar guides velvety, white

130

Bothalia 42,2 (2012)

speckled with purple or dark blue, often with larger

dark mark at base, often slightly scented of honey or

vanilla, outer tepals 20-31 mm long, claws pale mauve

often with darker central line and spotted purple-black,

10-12 mm long, velvety, limbs ± broadly lanceolate,

(9-)ll-18 X 10-14 mm, apex obtuse-apiculate, inner

tepals erect, hair-like, acute, 1.5-2. 5 mm long. Sta-

mens with filaments ± free, 3. 5-6.0 mm long; anthers

4.5-7.0(-8.0) mm long, pollen white (often red in west

of range, rarely pale blue). Ovary exserted from spathes,

6. 0-9. 5 mm long, flushed red; style vestigial, < 0.5 mm

long, branches ± 10 mm long; crests linear, erect or

arching inward, 6-9 mm long. Capsules narrowly ellip-

soid, 11-16 mm long, sometimes with thickened api-

cal rim. Seeds 1. 7-2.0 x ± 1 rnm, facets with spongy

edges and prominent spongy micropylar crest. Chromo-

some number 2n= \2 (Table 1). Flowering time: mostly

Sept. -mid-Oct. Figure 13.

Distribution: relatively widespread, Moraea ama-

bilis extends from the Olifants River Valley to the

Bokkeveld Escarpment and in an arc across the northern

edge of the Roggeveld Escarpment and south through

the Roggeveld and Klein Roggeveld to Worcester and

the western end of the Little Karoo (Figure 11). Plants

grow in light clay soils or in sandy loam. Locally in

the Bokkeveld Mtns and Gifberg, plants grow in shal-

low sandy ground on sandstone pavement where they

are usually dwarfed, we assume because of the shallow,

nutrient poor soil. In the west of its range plants flower

from late August to mid- September, but at higher eleva-

tions to the east, plants bloom from mid September into

October, sometimes as late as the end of that month.

Diagnosis: Moraea amabilis is recognized in the

Moraea tripetala complex by the virtually free filaments

(united for < 4 mm) and inner tepals reduced to hair-like

cusps up to 2.5 mm long, and by the distinctive mode

of vegetative reproduction in which the parent corm is

replaced annually by two new corms. Plants also often

have a cormlet in the leaf axil and as a result form clonal

colonies. This unusual characteristic is only evident if

the plants are carefully extracted from the ground (see

Figure 14A). The species is most likely to be confused

with M grandis, which also has virtually free filaments

but that species is larger in all features, notably the

ovary, 6-9 mm long in M amabilis vs. 10-12(-15) mm

in M. grandis, and anthers 4.5-7.0(-8.0) mm and up to

half again as long as the filaments vs. 9-11 mm long and

twice as long as the filaments in M. grandis. The corms

of M. amabilis have pale (rarely dark) brown, coarsely

fibrous tunics with prominent vertical elements in con-

trast to the finer tunic fibres in M grandis. Apart from

smaller flowers, M. amabilis often has white pollen in

contrast to the consistently bright red-orange pollen of

M. grandis (Table 2).

Capsules of Moraea amabilis, typically 11-16 mm

long, contain relatively small, sharply angular seeds, ±

2 mm at longest axis, with a prominent, pale, spongy

micropylar crest (Figure 14D). In contrast, capsules of

M. grandis are significantly larger, up to 19 mm long,

have a thickened apical ridge and the large, dark brown

seeds are up to 3 mm long (Figure 15D. A velvety to

pilose pubescence is universal on the leaves, stem and

spathes of western Karoo populations of M. amabi-

lis but those from the Bokkeveld Mtns, Gifberg and

Cedarberg-Olifants River Mtns usually have smooth

leaves although some notable exceptions include Gold-

blatt & Porter 13522 from Kransvlei and Levyns 10155

from Paleisheuwel. We place only moderate confidence

in the pubescence character as some other members of

the M. tripetala complex occasionally have hairy leaves

(notably M mutila), but we have seen no collections

of M. grandis with hairy leaves. On the dolerite flats

near Farm Keiskie, southeast of Calvinia, plants of M.

grandis {Goldblatt et al. 13366) were entirely hairless,

whereas M. amabilis growing nearby had pubescent

leaves, stems and sheathing leaves {Goldblatt et al.

13365). The possibility that M. amabilis and M grandis

hybridize in the Nieuwoudtville area where the two co-

occur is discussed briefly under the latter species.

Plants from near Worcester match Moraea amabi-

lis vegetatively, notably in their corm tunics and vel-

vety leaves and stems but have unusual, buff-yellow, or

dusty pink flowers (e.g. Goldblatt 4089). Plants from the

Rabiesberg, not far distant, also have velvety stems and

leaves and were described as having brown (we assume

buff-brown) flowers. These colour variants are unique

for the M. tripetala complex.

Moraea amabilis was described by Friedrich Diels

in 1910 based on plants he had collected in September

1900 when he lived in Calvinia. Goldblatt (1976b) erro-

neously designated Diels 626 as the holotype, overlook-

ing the second collection cited in the protologue, Diels

1169 (incidentally no longer extant). The so-called hol-

otype becomes the lectotype. Treated merely as a vari-

ant morph of M tripetala by Goldblatt (1976b), we now

regard M. amabilis as one the most distinctive members

of the M. tripetala complex in its habit of often produc-

ing two new corms in place of the single parent corm.

Flowers of the type, evidently somewhat shrunken,

have tepal claws 10-12 mm long and limbs ± 12-14 x

1 1 mm, an ovary ± 7 mm long, and the stamens have

filaments ±3.5 mm and anthers 5-6 mm long. Leaves

and corms are lacking, but as these were described they

were presumably present in the syntype, now lost, Diels

1169. The extant type material most closely matches the

morph from the sandstone pavement in the Bokkeveld

Mtns close to the edge of the escarpment.

Representative specimens

NORTHERN CAPE. — 3119 (Calvinia): between Oorlogskloof

and Papkuilsfontein, (-AC), Sept. 1939, Leipoldt 3059 (BOL),

Leipoldt 3002 (BOL, PRE); Oorlogskloof Nature Reserve, sandstone

pavement near entrance, burned last summer, (-AC), 13 Sept. 2010,

Goldblatt & Porter 13532 (MO, NBG, PRE); 0.5 km S of top of Van-

rhyn’s Pass, (-AC), 29 Aug. 1974 Goldblatt 2471 (MO; 15 km SSW

of Nieuwoudtville, Oorlogskloof Nature Reserve, (-AC), 31 Aug.

1995, Pretorius 259 (NBG, PRE); Lokenburg, rugged TMS, (-CA), 28

July 1956, Acocks 18900 (K, PRE); Farm Keiskie, dolerite flats SE of

Keiskie-se-Poort, plants hairy (co-blooming with M grandis), (-DB),

16 Sept. 2009, Goldblatt, Manning & Porter 13365 (MO, NBG). 3120

(Williston): ± 70 km SE of Calvinia to Middelpos on Blomfontein

road, (-CC), 26 Oct. 1976, Goldblatt 4381 (MO, PRE). 3220 (Suth-

erland): Roggeveld, Soekop, Rooipunt Camp, (-AA), 17 Sept. 2004,

Rosch 253 (NBG); Roggeveld Plateau, Farm Blesfontein, sandy flats,

(-AD), 4 Nov. 2011, Goldblatt & Porter 13720 (MO, NBG); Smous-

hoogte, slopes above road (visited by Rediviva macgregorii), (-DC),

17 Sept. 1993, Steiner 2710 (MO, NBG); top of Verlaten Kloof, (-

DC), 28 Oct. 1982, Bayer 3276 (NBG); Moordenaars Karoo on road

to farm Meintjiesplaas, shaded S slope above stream, (-DC), 30 Sept.

Bothalia 42,2 (2012)

131

FIGURE 13. — Moraea amabilis, Goldblatt & Porter 13461. A, flowering stems and corm; B, inner perianth whorl plus filaments and style; C,

capsules; D, seeds. Scale bar: A, C, 10 mm; B, D, 2 mm. Artist: John Manning.

FIGURE 14.— A/oraea grandis, Goldhialt & Manning 13658. A, flowering stems and corm; B, inner perianth whorl plus filaments and style; C,

capsules; D, seeds. Scale bar: A, C, 10 mm; B, D, 2 mm. Artist: John Manning.

Bothalia 42,2 (2012)

133

TABLE 2, — Comparison of taxonomically important features in Moraea amabilis and M grandis. Filaments are virtually free or basally united for

less than 0.4 mm in all specimens of both species examined.

2004, Goldblatt & Porter 12656 (MO); Klein Roggeveld, (-DC), Oct.

1920, Marloth 9595 (PRE), 9602 (PRE).

WESTERN CAPE. — 3118 (Vanrhynsdorp): Gifberg, plateau, (-

DB), 23 Aug. 1984, Goldblatt 7230 (MO), 13 Sept. 2010 (fr.), Gold-

blatt & Porter 13524 (MO, NBG), 23 Jun. 1913, Phillips 7524 (K);

Bulhoek, (-BD), 2 Aug. 1896, Schlechter 8382 (MO, PRE); top of

Nardouw Pass, (-BD), 13 Aug. 1976, Goldblatt 3853 (MO). 3218

(Clanwilliam): Clanwilliam, at tumofif to Boskloof, (-BB), 29 Aug.

1974, Goldblatt 2457 (MO, PRE), 11 Sept. 2010 (ff.), Goldblatt &

Porter 13522A (MO, NBG); W slopes of Pakhuis Pass, 10 km from

Clanwilliam, (-BB), [?date], Nordenstam & Lundgren 1325 (MO, S);

sandy slopes near Paleisheuwel, (-BC), 5 Sept. 1954, Levyns 10155

(BOL). 3219 (Wuppertal): Klipfonteinrand, (-AA), 20 Aug. 1974,

Nordenstam & Lundgren 1347 (MO, S). 3220 (Sutherland): Koedoes

Mtns, rocky gully above stream, (-CC), 14 Sept. 2009, Goldblatt,

Manning & Porter 13327 (NBG, MO); 70 km S of Sutherland to

Matjiesfontein, (-DC), 30 Oct. 1974 (late fl. & fr.), Goldblatt 3222

(MO); N of Farms Fortuin and Nuwerus at Komsberg turnoff from

Sutherland road (co-blooming with M cuspidata), (-DC), 28 Sept.

2009, Goldblatt & Porter 13461 (MO). 3319 (Worcester): Rabiesberg,

Montagu, (flowers brown), ex hort. Kirstenbosch, (-CA), July 1940,

Van der Gaast s.n. (Nat. Bot. Gard. 2673/35 in NBG); Rabiesberg,

lower slopes, flowers pink, (-CA), 27 Sept. 1935, Lewis s.n. (BOL

21733, K); foot of Fonteintjiesberg, Farm Onse Rug, clay soil, flow-

ers pink or buff, (-CB), 12 Sept. 1976, Goldblatt 4089 (K, MO, PRE).

3320 (Montagu): Memorial Hill, rocky slope above cemetery, (-AB),

29 Sept. 2009, Goldblatt & Porter 13467 (MO); S-facing bank of

stream opposite Pietermeintjies Siding, (-AC), 23 Sept. 1976, Gold-

blatt 4175 (MO, PRE); Op-de-Tradouw, Barrydale to Montagu, (-DC),

Goldblatt 4185 (MO).

9. Moraea grandis Goldblatt & J.C. Manning, sp.

nov.

TYPE. — ^Northern Cape, 3119 (Calvinia): Wild

Flower Reserve at Nieuwoudtville, edge of shrubs near

top of dolerite slope, scented of vanilla, (-AC), 22 Sept.

2011, Goldblatt & Manning 13658 (NBG, holo.; MO,

PRE, iso.).

Plants 200-400 mm high, usually in small colonies.

Corm mostly 10-15 mm diam, with tunics of light

(rarely dark brown), relatively fine, firm or soft fibres,

rarely accumulating in a thick mass; often producing

two new conns to replace parent corm. Stem glabrous,

rarely velvety to pilose, simple or 1 (2)-branched, base

sheathed by dry light brown cataphylls, these usually

persisting as collar of fine fibres around base; usually

producing one or more cormlets in leaf or cataphyll

axils. Foliage leaf solitary, shallowly channelled, usu-

ally exceeding stem and up to twice as long, 3. 5-5. 5

mm wide, glabrous; cauline sheathing leaves green or

becoming dry from tips, long-attenuate, 55-70 mm

long. Rhipidia several flowered; spathes green with dry,

attenuate tips, glabrous, inner 55-80 mm long, outer ±

half as long as inner. Flowers pale blue to pale violet

or light purple, nectar guides small, triangular, yellow

(rarely cream) on a white background and edged dark

blue, short-velvety (rarely glabrous), scented of vanilla.

outer tepals 30-40 mm long, claws 12-18 mm long,

velvety, limbs ± orbicular, widest above middle, usu-

ally ± as wide as long, 15-20(-23) x 15-20 mm, plane,

obtuse-apiculate, spreading at 45-60°, inner tepals hair-

like, erect, 1.5-3. 5 mm long. Stamens with filaments

± free or united < 0.2 mm, 4. 0-5. 5 mm long; anthers

8-11 mm long, usually not reaching stigma lobes, pol-

len orange-red. Ovary (10-)12-15 mm long, slightly

narrowed below apex; style vestigial, < 0.4 mm long,

branches ± 15 mm, crests linear, 8-12 mm long. Cap-

sules 16-19 mm long, narrowly ovoid-oblong with

thickened rim. Seeds relatively large, 2-3 x 1.8-2. 3 mm,

facets rounded, angles with wing-like ridges. Chromo-

some number 2n= \2 (Table 1). Flowering time: mainly

late Aug.-late Sept. Figure 14.

Distribution: restricted to the northern end of the

western (winter rainfall) Karoo, Moraea grandis extends

from the Langberg west of Loeriesfontein through the

higher country between Loeriesfontein and Calvinia

and along the Bokkeveld Plateau to the northern end of

the Roggeveld Escarpment (Figure 15). Plants favour

heavier soils and are most often found in heavy red

clay among dolerite rocks but also grow on lighter soils

derived from tillite and shale.

Diagnosis: the most striking of the species of the

Moraea tripetala complex, M. grandis has relatively

large corms up to 15 mm diam, with tunics usually of

fine, soft (rarely firm) fibres, and the largest flowers in

the alliance. The outer tepals are 30^0 mm long (Table

2) and have ± orbicular limbs 18-23 x 15-20 mm, thus

almost as wide as long, and have unusually small nec-

tar guides, usually yellow (sometimes cream) on a

white background, minutely spotted with dark blue, and

usually edged with a darker colour than the rest of the

tepal limb. The limbs are also widest in the upper third

134

Bothalia 42,2 (2012)

and usually have an abrupt, comma-like tip. The flow-

ers have a pleasant, vanilla-like odour. Like its relative,

M amabilis, it typically grows in colonies resulting

from vegetative reproduction by axillary cormlets and

the sometimes replacement of the parent corm by two

daughter conns. It also resembles M. amabilis in having

virtually free filaments (united for < 0.2 mm), but the

anthers, 8-1 1 mm long, are at least twice as long as the

filaments and sometimes 2.5 x as long, easily the largest

in the M tripetala complex. The pollen is consistently

bright orange-red, not unusual for the entire group, and

contrasting with the white pollen in most western Karoo

populations of M. amabilis, which has smaller anthers

4. 5-7.0 mm long and ± as long as or up to 1.5 x as long

as the filaments.

Moraea grandis sometimes grows in close proxim-

ity to M. amabilis, and the two can be seen flowering

together on the farm Keiskie south of Calvinia. Diffi-

culty in distinguishing the two in the immediate vicinity

of Nieuwoudtville is probably the result of hybridization

and introgression between the two species, which share

the same large-bodied bees as pollinators.

Moraeq grandis has been illustrated in several wild

flower volumes notably in the Nieuwoudtville, Bokkeveld

Plateau and Bantam wildflower guide (Manning &

Goldblatt 1997; 69) and The color encyclopedia of Cape

bulbs (Manning et al. 2002: 19, 307). The photographs

show the distinctive, broad outer tepal limbs oriented

almost vertically in fully open flowers.

l'

Representative specimens

NORTHERN CAPE. — 3018 (Kamiesberg): Langberg, wet gully

on shale, (-DB), 5 Sept. 2006, Goldblatt & Porter 12765 (MO, NBG).

3019 (Loeriesfontein): slopes of Kubiskou Mtn, clay ground, (-CC),

6 Sept. 2006, Goldblatt & Porter 12783 (MO); ± 8 km W of Loer-

iesfontein, slate hills, (-CD), 12 Sept. 1976, Thompson 2883 (NBG,

PRE). 3119 (Calvinia): Nieuwoudtville, Farm Glenlyon, (-AC), 2

Sept. 1959, Hardy 73 (PRE); 3 rniles [4.5 km], W of Nieuwoudtville,

(-AC), 28 Aug. 1950, Lewis 2274 (PRE); Glen Lyon; 2 mi. S of Nieu-

woudtville, red clay flats. Flowers blue. Fairly frequent in between

bulbinellas and rhenosterbos, (-AC), 11 Sept. 1974, Oliver & Mauve

42 (K, PRE); foot of Vanrhyns Pass, (-AC), 16 Oct. 1974 (ff.), Gold-

blatt 3098 (MO); Calvinia District, 8 km on Rietfontein road, W of

Calvinia-Loeriesfontein road, (-AD), 25 August 1976, Goldblatt 3948

(MO, PRE); Langfontein ± 3 km to Kotzeskolk, (-BB), 2 Sept. 1986,

Burger & Louw 184 (NBG); Farm Perdekraal, S slope on dolerite, (-

BC), 12 Sept. 1981, Goldblatt 6247 (MO); Hantamsberg, upper slopes

NW of Calvinia, (-BC), 16 Sept. 1980, Goldblatt 1980 (MO); road to

Toring, 4.1. km N of Loeriesfontein-Calvinia road, dolerite slope with

heavy red clay, (-BD), 21 Sept. 2011, Goldblatt & Manning 13655

(MO, NBG); Bloukrans Pass, near summit in wet gully, (-DA), 31

Aug. 1981, Snijman 622 (NBG, PRE); Farm Keiskie, dolerite flats SE

of Keiskie-se-Poort (co-blooming with M. amabilis), (-DB), 16 Sept.

2009, Goldblatt, Manning & Porter 13366 (MO, NBG).

ACKNOWLEDGEMENTS

We thank Elizabeth Parker and Lendon Porter for

their assistance and companionship in the field; Koos

Claassens of Jacobsbaai for help in investigating the sta-

tus of M. hainebachiana; Mary Stiffler, Research Librar-

ian, Missouri Botanical Garden, for help with literature

searches; Roy Gereau, Missouri Botanical Garden, for

assistance with nomenclatural questions; and Clare

Archer for lists of exsiccatae from the PRE collection.

Rhoda McMaster provided the initial stimulus for this

study by questioning the range of seed types in plants

then treated as Moraea tripetala. Collecting permits

were provided by the Nature Conservation authorities

of Northern Cape and Western Cape, South Africa. Field

work was funded in part by the Mellon Foundation and

the B.A. Krukoff Fund, Missouri Botanical Garden.

We acknowledge the generous donation of the Haine-

bach family of Camps Bay, Cape Town, for conservation

of the Cape Flora. Their contribution is remembered in

the name Moraea hainebachiana. We are also delighted

to recognize the valuable support towards our research

provided by Ms Naoko Ogama, Miyagi prefecture,

Japan, by commemorating her in the name Moraea oga-

mana.

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Press, Calcutta.

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Bothalia 42,2: 137- 145 (2012)

New species and subspecies of Babiana, Hesperantha^ and Ixia

(Iridaceae: Crocoideae) from southern Africa; range extensions and

morphological and nomenclatural notes on Babiana and Geissorhiza

P. GOLDBLATT* and J.C. MANNING"

Keywords: Babiana Ker GawL, Geissorhiza Ker Gawl., Hesperantha Ker Gawl., Iridaceae, Ixia L., new species, southern Africa, taxonomy

ABSTRACT

Babiana rivulicola from stream banks in the Kamiesberg in Namaqualand and terete-leaved Ixia teretifolia from the

Roggeveld, both in Northern Cape, are new species of these two largely winter-rainfall region genera. Late-flowering popula-

tions of Hesperantha radiata with crowded spikes of smaller flowers are segregated from the typical form as subsp. caricina.

We also document the first record of B. gariepensis from Namibia, correct the authority for B. purpurea Ker Gawl., discuss

morphologically aberrant populations of B. tubiflora from Saldanha, provide an expanded description for B. lapeirousiodes

based on the second and only precisely localized collection of this rare Namaqualand species, and expand the circumscription

of Geissorhiza demissa to accommodate a new record from the Kamiesberg, including revised couplets to the existing key

to the species.

INTRODUCTION

Crocoideae, with ± 1 080 species, are the largest of

the six subfamilies of Iridaceae, which now include

over 2 200 species worldwide. Largely sub-Saharan

African, subfamily Crocoideae is centred in south-

ern Africa, where some 980 species occur, almost 850

of them restricted to the southwest of the subcontinent

(the Greater Cape Floristic Region). The richness of the

Cape flora is well established (e.g. Goldblatt & Man-

ning 2000) and although the region has been explored

botanically for over 250 years, novelties are still regu-

larly recorded. Some are completely new discoveries

but many others have reposed in herbaria, where they

are recognized only after critical revisions of particular

groups are undertaken. In Iridaceae alone, 97 species

have been added to the family for the Cape flora over the

past decade.

Here we describe one new species each of Babiana

Ker Gawl. (now 93 spp.) and Ixia L. (now ± 90 spp.).

Both genera are members of tribe Ixieae Dumort. (1822)

as circumscribed by Goldblatt et al. (2006) and, apart

from Ixia, have been revised or thoroughly reviewed in

the past 25 years (Goldblatt 2003; Goldblatt & Man-

ning 2007a, 2010). In Hesperantha (82 spp.), we also

describe a new subsp. of Hesperantha radiata (L.f)

Ker Gawl and include a significant range extension into

Namibia for B. gariepensis Goldblatt & J.C.Manning,

morphological notes for variant populations of B. tubi-

flora (L.f.) Ker Gawl. and Geissorhiza demissa Gold-

blatt & J.C.Manning, nomenclatural notes for B. bainesii

Baker, and a correction to the author citation for B. pur-

purea Ker Gawl.

■ B.A. Krukoff Curator of African Botany, Missouri Botanical Garden,

RO. Box 299, St. Louis, Missouri 63166, USA. E-mail: peter.gold-

blatt@mobot.org.

” Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, Claremont 7735, South Africa / Research Centre for

Plant Growth and Development, School of Life Sciences, University of

KwaZulu-Natal, Pietermaritzburg, Private Bag XOl, Scottsville 3209,

South Africa. E-mail: J.Manning@sanbi.org.za.

MS. received: 2011-12-22.

MATERIALS AND METHODS

We examined all relevant collections at BOL, NBG,

PRE, and SAM, the primary southern African herbaria

(acronyms after Holmgren et al 1990).

SYSTEMATICS

New taxa

1. Babiana Ker Gawd.: Lewis’s (1959) monograph

of Babiana recognized 61 species, including one from

the Indian Ocean island of Socotra, and set the stage

for later studies of the genus, not least by stabilizing its

complex synonymy. Antholyza L. (including Anaclanthe

N.E.Br.), with two species, was added to the genus by

Goldblatt (1990). In the recent revision of Babiana

(Goldblatt & Manning 2007a) we recognized 88 species,

excluding the Socotran B. socotrana Hook.f, which is

now the new genus Cyanixia Goldblatt & J.C.Manning

(Goldblatt et al 2004). Subsequent exploration of west-

ern South Africa yielded an additional four new spe-

cies (Goldblatt et al 2008; Goldblatt & Manning 2010).

The discovery by Cape Town biologist Nick Helme of

yet another new species, described here as B. rivulicola,

in a relatively well collected part of the Kamiesberg in

central Namaqualand in 2009 is thus particularly surpris-

ing. Babiana now has 93 species, but additional taxa are

likely to be recognized as the results of the molecular

phylogenetic study (Schnitzler et al 2011) are analyzed

from a systematic perspective.

Babiana rivulicola Goldblatt & J.C.Manning, sp.

nov.

TYPE. — ^Northern Cape, 3018 (Kamiesberg): south-

ern Kamiesberg, Langkloof, ± 20 km from Doringkraal

turnoff from N7, Farm Nartjiesdam, in rocky bed of per-

ennial stream, (-AD), 22 Sept. 2010, Goldblatt & Porter

13572 (NBG, holo.; K, MO, PRE, iso.).

Plants to 300 mm high, excluding longer leaves,

growing in dense clumps; stem erect, simple or

138

1-3-branched, angled. Corm wedged in crevices in

rock, unknown. Leaves mostly 6 or 7, linear, 250^50

X 5-10 mm, exceeding stem, firm but arching toward

ground, shallowly pleated to almost flat, glabrous.

Spike flexed outward with flowers initially in two ranks,

mostly 6-9-flowered; bracts green below, dry and light

brown toward tips, outer mostly up to 35 mm long but

lowermost much longer, attenuate, inner ± 2/3 as long

as outer, divided apically, attenuate, 2-keeled below.

Flowers zygomorphic, pale blue to blue-mauve, paler

in throat, lower 3 tepals with central, white, lanceolate

blotch and darker violet marking toward base, darkly

lined on abaxial side of throat, lightly sweet-scented;

perianth tube funnel-shaped, 42-44 mm long, lower part

cylindric, upper ± 10 mm expanded into wide throat;

tepals lanceolate, unequal, dorsal ± 30 x 8 mm, lower

three tepals united with upper laterals for ± 3 mm, arch-

ing outward, 25 x 6 mm. Stamens unilateral; filaments

straight, suberect, ± 15 mm long, included in floral cup,

reaching almost to middle of dorsal tepal; anthers ± 5

mm long, pale mauve, pollen white. Ovary glabrous,

style dividing opposite or slightly beyond anther tips,

style branches ± 5 mm long, recurved distally. Capsules

and seeds not known. Flowering mid- to late Oct. Figure

1.

Bothalia 42,2 (2012)

Eponymy. from the Latin, rivulieola, growing in a riv-

erine habitat.

Distribution and ecology, so far Babiana rivulieola

is known only from the southern Kamiesberg in North-

ern Cape, where it is restricted to the perennial stream

that runs through the Langkloof, between Farms Karas

(Welkom) and Doringkraal (Figure 2). Plants grow in

cracks in smooth granite along the edges of the stream

above the waterline. Plants are regularly inundated

after heavy rains but the corms are wedged in crev-

ices in bedrock and secure from being washed away. It

proved impossible to extract corms for examination and

they remain unknown. The species was first collected

in October 2009 by Cape Town biologist N.A. Helme.

Despite its narrow range, we see no current threat to B.

rivulieola and suggest a conservation status of (LC),

least concern, using the definitions and terminology of

Raimondo et al. (2009).

Diagnosis and relationships: Babiana rivulieola most

closely resembles a second, more widespread Namaqua-

land endemic, B. dregei Baker, and both species share

glabrous leaves, stem and bracts, an unusual feature in

a genus where pubescence of some kind on vegetative

FIGURE 1 . — Babiana rivulieola,

Goldblatt & Porter 13572

(NBG). A, portion of flower-

ing plant; B, outer (left) and

inner (right) bract; C, half

flower. Scale bar: 10 mm. Art-

ist: J.C. Manning.

Bothalia 42,2 (2012)

139

FIGURE 2. — Distribution of Babiana rivulicola, o; and Ixia tereti-

folia, •.

organs is almost universal. The leaves of B. dregei are

characteristically relatively broadly lanceolate and rigid

with sharply pungent tips and the stem is very short and

concealed among the leaves. In B. rivulicola the stem

reaches up to 250 mm high and the linear leaves are

longer and relatively softly textured, with acute but not

pungent tips. The flowers in the two species are broadly

similar, ± fuimel-shaped with the lower two thirds of

the perianth tube cylindric, but B. dregei has a ± cylin-

dric perianth tube (45-)50-65(-70) mm long, curved

just below the apex, unlike the shorter, funnel-shaped

perianth tube of B. rivulicola, 42-44 mm long. Addi-

tional floral differences include the longer anthers, ± 8

mm long and shorter style branches, 3-4 mm long in

B. dregei vs. anthers ± 5 mm long and style branches ± 5

mm long in B. rivulicola.

When collecting Babiana rivulicola in the second

half of September we noted plants of B. dregei on adja-

cent slopes well past flowering and with developing cap-

sules. Although sympatric in distribution, the two taxa

are isolated by flowering time and ecology, and thus

allotopic (Wiley 1981).

Additional specimen

NORTHERN CAPE. — 3018 (Kamiesberg): southern Kamiesberg,

Langkloof, ± 1 km S of Farm Nartjiesdam, in rocks in stream, (-AD),

10 Oct. 2009, Helme 6110 (NBG).

2. Hesperantha Ker Gawl.\ the southern and tropi-

cal African Hesperantha with 82 species (Goldblatt

2003; Goldblatt & Manning 2007b), is distinguished

in Crocoideae by the style dividing at the mouth of the

perianth tube (rarely within the tube) into long, spread-

ing, relatively long, more-or-less straight, laxly spread-

ing style branches and, with a few exceptions, by hard,

woody corm tunics (Goldblatt & Maiming 2008). In

the most widespread species of the genus, H. radiata,

Goldblatt (2003) drew attention to collections of small-

flowered plants from the Cape Peninsula and surrounding

hills, as far north as Piketberg and as far east as Hermanns

in the southwestern comer of the southern African winter-

rainfall zone, which were then included in the more wide-

spread and common form which extends from Namaqua-

land to Swaziland. These small-flowered plants have

unusually crowded spikes with shortly imbricate bracts,

and the uppermost leaf is always entirely sheathing and

reaches almost to the base of the spike. The flowers have

tepals 7-12 mm long, a tube 6-10 mm long, anthers 4(-5)

mm long, and the stems have a weakly developed neck

of fine fibers around the base (Table 1). Including this

morph in H. radiata, of which typical larger flowered

populations occur in the same area, seems counter to the

concept of biological species. Elsewhere, the distinc-

tion between the two forms is not always entirely clear,

which argues against treating them as distinct species. As

Goldblatt (2003) also pointed out, it is especially nota-

ble that the small-flowered plants bloom later than typi-

cal H. radiata and are sometimes sympatric but not co-

blooming [allotopic (Wiley 1981)], as for example, Oliver

4332 (NBG) collected on 21 Aug. 1973 and 4756 ( NBG)

collected on 17 Oct. 1973, both in the hills at Langver-

wacht near Stellenbosch. The August-flowering plants are

typical H. radiata and those flowering in October are the

small-flowered morph with crowded spikes. Specimens

of this morph collected by C.F. Ecklon and C.L. Zeyher

in the early 19th century are annotated H. setacea Eckl.

(e.g., Ecklon & Zeyher Irid. 233 89.9), the name a nomen

nudum and thus invalid (Ecklon 1827), while some sheets

of the small-flowered plants at the Kew Herbarium are

annotated H. tenuifolia. This is R.A. Salisbury’s (1812)

name at species rank for H. radiata van y caricina of Cur-

tis’s Botanical Magazine t. 790 (Ker Gawler 1804). Salis-

bury’s epithet alludes to the characteristic narrow leaves,

also found in some populations of larger-flowered plants

that correspond to the type of H. radiata.

We propose to recognize late-blooming plants

with smaller flowers as a separate subspecies, in line

with our treatment of late-flowering Moraea tripetala

subsp. jacquiniana (Schltr. ex G.J.Lewis) Goldblatt &

J.C.Manning, which is sympatric with early flowering

subsp. tripetala (Goldblatt & Manning 2012a).

TABLE 1 . — Comparison of flowers of Hesperantha radiata subsp. car-

icina and subsp. radiata. Only plants with well pressed, fully

open flowers were measured.

Hesperantha radiata subsp. caricina (Ker Gawl.)

Goldblatt & Manning stat. nov. H. radiata var. y cari-

cina Ker Gawl.: t. 790 (1804). H. tenuifolia Salisb.: 321

(1812), nom. nov. pro H. radiata var. y caricina Ker

Gawl. (1804). H. caricina (Ker Gawl.) Klatt: 395 (1882),

nom. illegit. superfl. pro H. tenuifolia Salisb. (1812). H.

angustifolia Loudon: 91 (1841), nom. illegit. superfl. pro

H. tenuifolia Salisb. (1812). Type: South Africa, without

precise locality, illustration in Curtis’s Botanical Maga-

zine 21: t. 790(1804).

Plants like subsp. radiata but corms often with a col-

lar of fibres around base. Leaves 1-2 mm wide, upper-

most leaf entirely sheathing, reaching close to base of

spike. Spike crowded with outer bracts 10-13 mm long.

140

Bothalia 42,2 (2012)

as long as or longer than next intemode and thus imbri-

cate. Flowers cream to dull yellow, outer tepals reddish

on reverse; perianth tube 6-10 mm long; tepals 7-12 x

2. 5-3. 5 mm. Stamen filaments 4—5 mm long; anthers

4(-5) mm long. Style branches mostly 5-6 mm long.

Flowering time: mostly late Sept, and Oct., but until Dec.

at higher elevations.

Diagnosis: subsp. caricina is usually recognized by

the crowded spike of relatively small flowers, the tepals

7-12 X 2. 5-3. 5 mm and perianth tube 6-10 mm long,

and shortly overlapping bracts 10-13 mm long. Plants

are typically late flowering, in September and October,

but there are collections from higher elevations that

bloom as late as December. Populations of typical H.

radiata occur throughout the range of subsp. caricina

(from Piketberg to Caledon and Hermanns) but invari-

ably flower earlier, from late July to mid-September,

and have larger flowers, with tepals 13-15(-17) x 3-5

mm, perianth tube 8-14 mm long, and anthers mostly 4

mm long (Table 1 ). The leaves of subsp. caricina are also

somewhat narrower than in subsp. radiata,\-2 mm wide,

and often dry at flowering time vs. 2.0-3. 5 mm wide and

green at flowering.

Representative specimens

WESTERN CAPE. — 3318 (Cape Town): Riebeek Kasteel, Farm

Remhoogte, (-BD), 24 Oct. 1968, Marsh 1034 (NBG); Table Mtn, top

of Oudekraal Ravine, (-CD), Nov. 1972, McKinnon s.n. (NBG); Rose-

bank, Cape Town, Nov. [without year], H. Bolus 3769 (BOL, K, NBG,

PRE, Z); Langverwacht above Kuils River, (-DC), 17 Oct. 1973,

Oliver 4756 (K, NBG, PRE); Stellenbosch flats, (-DD), 30 Sept. 1958,

Perold 7 (NBG); Sept. 1964, Holzapfel 10 (NBG); Blauwklip, (-DD),

24 Oct. 1928, Gillelt s.n. (NBG); Uitkyk, (-DD), Oct. 1982, Gillett s.n.

(NBG). 3418 (Simonstown): flats between Gordon’s Bay and Strand

on Disa Street, (-BB), 29 Sept. 2001, Goldblatt & Ndnni 11944 (MO,

NBG). 3419 (Caledon): hills near Farm Die Vlei E of Bot River, (-

AA), 30 Sept. 2000, Goldblatt & Ndnni 11944 (MO); western outskirts

of Caledon, (-AB), 29 Sept. 2001, Goldblatt & Ndnni 11942 (MO);

Femkioof Nature Reserve, Hermanus, (-AD), 18 Oct. 1979, Orchard

4485 (MO). Without precise locality: Piketberg, renoster hills, 9 Oct.

1922, Marloth 11484 (NBG).

3. Ixia L.; restricted to South Africa and to the win-

ter rainfall zone and adjacent areas, Ixia comprises an

estimated 90 species in four sections. Sects. Hyalis (18

spp.), Morphixia (now 32 spp.), and Dichone (now 17

spp.) have recently been revised (Goldblatt & Manning

2011, 2012b) and we are currently revising the system-

atics and taxonomy of sect. Ixia (± 25 spp.). Here we

describe a species in sect. Morphixia that we discovered

on the edge of the Roggeveld Escarpment, in September

2011, and later collected in fruit in November.

Ixia teretifolia Goldblatt & J.C. Manning, sp. nov.

TYPE. — Northern Cape, 3220 (Sutherland): Rog-

geveld Escarpment, Farm Blesfontein, in dense tall grass

tufts below sandstone cliffs at escarpment edge, (-AD),

25 Sept. 2011, Goldblatt & Manning 13671 (NBG,

holo., K, MO, PRE, iso.).

Plants mostly 1.2-1. 5 m high. Corm subglobose,

12-18 mm diam., tunics of relatively fine fibres. Stem

erect in flower, nodding in fruit; sheathed below by

prominent, pale, dry-membranous, often irregularly tom

cataphylls accumulating with age in a dense mass; usu-

ally with 3-5 short branches up to 4 mm long, branches

subtended by pale, dry, attenuate scale-like bracts and

prophylls 2-A mm long. Leaves 3, lower 2 with long

terete blades ± half as long as stem, mostly 1. 5-2.0 mm

diam., flrm-textured without evident veins when alive;

uppermost leaf entirely sheathing or almost so, reach-

ing to base of spike or to lowermost branches. Main

spike ( 1)2(3 )-flowered, lateral spikes 1- or 2(3)-flow-

ered; bracts silvery membranous, translucent with nar-

row, dark purple veins, ± 7 mm long, outer 3 veined

and 3-toothed or 3-lobed; inner 2-veined and shallowly

forked at apex. Flowers upright to half nodding, pale

blue but pale green in throat, faintly rose-scented, with

bright yellow anthers prominently displayed; perianth

tube broadly funnel-shaped, ± 5 mm long, narrow part ±

2 mm long; tepals spreading, fully patent distally, ovate,

± 12 X 6. 0-6. 5 mm, outer tepals slightly wider than

inner. Stamens with filaments ± 6 mm long, exserted ± 4

mm from tube; anthers 4-5 mm long, ± parallel through-

out anthesis, yellow; pollen yellow. Style dividing oppo-

site lower third of anthers, branches ± 3 mm long, slen-

der, extending between anthers. Capsules globose, ± 5

mm diam., showing outline of seeds. Seeds subglobose,

± 2.0 X 1.6 mm, light yellow-brown, (5)6 per locule.

Flowering time: mostly mid-Sept.-mid-Oct., possibly

later. Figure 3.

Eponymy: from the Latin, teres, round in section,

folia, leaf.

Distribution and ecology: so far known from just

one site, Ixia teretifolia grows at the base of sandstone

cliffs of the Beaufort Series at the edge of the Roggeveld

Escarpment west of Sutherland on the Farm Blesfontein

(Figure 2). Plants grow among broken rocks, nested in

dense tufts of the grass Tenaxia (Merxmuellera) stricta

(Schrad.) N.P.Barker & H.P.Linder. Extensive searching

on the escarpment itself revealed no sign of the species,

but the white -flowered morph of the related 7. margin-

ata Salisb. ex G.J. Lewis was relatively common there,

growing in mountain renosterveld vegetation together

with other geophytes, including 7 trifolia G.J.Lewis,

Geissorhiza heterostyla L.Bolus, the deep pink-flowered

morph of Hesperantha pilosa (L.f ) Ker Gawl. (all Iri-

daceae) and Bulbinella elegans P.L.Perry (Asphode-

laceae). We estimated that the population of 7 teretifo-

lia consisted of about 70 mature, flowering individuals

along a 200 m stretch of the escarpment edge. Although

we found no plants to the south of our site on the adja-

cent farm Boplaas, we suspect the range of 7. teretifo-

lia may be more extensive further north toward Ouberg

Pass, where we recommend further exploration to estab-

lish its complete range.

Diagnosis and relationships: the short, widely fun-

nel-shaped perianth tube, ± 5 mm long, distally spread-

ing tepals, partly exserted filaments with prominently

displayed anthers, and the several short lateral branch-

lets place 7 teretifolia in the informal series Marginatae

of Ixia sect. Morphixia (Goldblatt & Manning 2011).

Its tall stature and habit are shared in the section with

7 alata Goldblatt & J.C. Manning, but the several very

short branches, each with 1 or 2(3) flowers differ from

that species, and recall particularly 7 linearifolia Gold-

blatt & J.C. Manning. The centric leaves lacking visible

veins are unique in sect. Morphixia and unusual in the

genus, almost all species of which have plane, isobilat-

eral leaves, apart from 7 linearifolia, which has linear.

Bothalia 42,2 (2012)

141

FIGURE 3. — Ixia teretifolia, Gold-

blatt & Manning 13671

(NBG). A, portion of flower-

ing plant; B, T/S leaf blade;

C, leaves and flowering stem.

Scale bar; A-C, 10 mm; B, 1

mm. Artist: J.C. Manning.

142

Bothalia 42,2 (2012)

fleshy leaves with the margins raised at right-angles to

the blade surface. Except for their slightly smaller size,

the flowers of /. teretifolia differ very little from other

members of series Marginatae, all of which have a short,

funnel-shaped perianth tube, distally outspread tepals

and fully exserted anthers.

Internally, the leaf margins in I. teretifolia have paired

strands of sclerenchyma separated by parenchyma tissue

(Figure 3B). As in other species of Ixia the sclerenchyna

strands are not associated with a marginal vein ((Rudall

& Goldblatt 1991); Rudall’s (1995) statement to the con-

trary for Ixia is an error), and there are just two opposed

major vein pairs in the leaf in a lateral position, well

separated internally by mesophyll. The anatomy of I.

teretifolia does not conform to that of a truly centric (or

terete) leaf as it is not radially symmetric, e.g. as seen

in terete-leaved species of Bobartia (Rudall 1995), and

the leaves are in fact internally bilaterally symmetric.

Related I. linearifolia has fairly typical leaf anatomy for

Ixia with a wide strand of sclerenchyma extending the

width of the thickened, but slightly winged margins,

themselves unusual in Ixia.

Additional specimen

NORTHERN CAPE. — 3220 (Sutherland): Roggeveld Escarpment,

Farm Blesfontein, in dense tall grass tufts below sandstone cliffs at

escarpment edge, (-AD), 4 Nov. 2011 (ft.), Goldblatt & Porter 13723

(MO, NBG).

Morphological and nomenclatural notes and a

range extension

1. Babiana tubiflora (L.f) Ker GawL: a population

of a Babiana species on the inland side of high sand

dunes north of Jacobsbaai on the Saldanha Peninsula is

somewhat puzzling in its unusual flowers. These plants

are currently included in the widespread Babiana tubi-

flora, which extends along the coast of Western Cape

from Lambert’s Bay in the north to Still Bay in the east

(Goldblatt & Manning 2007a), thus surrounding the

Jacobsbaai population. We were unable to distinguish

the Jacobsbaai plants vegetatively from specimens of B.

tubiflora collected nearby at Langebaanweg when com-

pared side-by-side, but the flowers are strikingly differ-

ent. Largely a coastal species, B. tubiflora rarely extends

more than 20 km inland, and has zygomorphic flow-

ers with a white to cream-coloured perianth, the lower

tepals often with a red streak on the limbs of the lower

three tepals. The tepals are narrowed into an ascending

claw-like lower portion with wider, spreading limbs,

the dorsal tepal 18-23 x 5-7 mm and the lower tepals

15-19 X 3-4 mm. The unilateral stamens have filaments

13-16 mm long and anthers 4-5 mm long. In contrast

(Table 2), the tepals of the Jacobsbaai plants are ovate

without a narrowed claw, the dorsal tepal is 12 x 6 mm,

and the lower three tepals are ± 10 x 4 mm {Goldblatt

& Porter I35I2 MO, NBG, 9 Sept. 2010; Goldblatt &

Porter 13572, MO, NBG, 23 Sept. 2010). The stamens

have filaments ± 6 mm long (< half the length of those

of B. tubiflora as currently circumscribed) and anthers

± 3 mm long. The relatively short style divides opposite

the base to lower third of the anthers into branches ±2.5

mm long, whereas that in typical B. tubiflora divides

between the middle and slightly beyond the anther tips

and the style branches are 3-4 mm long. We note that

the Jacobsbaai B. tubiflora population was sympatric

and co-blooming with closely related but larger-flowered

B. tubulosa (Burm.f.) Ker Gawl. {Goldblatt & Porter

13566, MO, NBG, PRE, 22 Sept. 2010).

We remain uncertain whether to regard the Jacobsbaai

plants as a separate taxon or merely a trivial mutant race

of no taxonomic significance. Until more is known about

this distinctive population we simply report its exist-

ence and defer a decision about its status pending further

study.

2. Babiana purpurea Ker Gawl. : Ixia purpurea Jacq.

was based on a plate in his leones plantarum rariorum

vol. 2: t. 286 (Jacquin 1793), but with the description

published earlier (Jacquin 1791). The name was later

transferred to Babiana by Ker Gawler (1807a) and has

been cited until now as B. purpurea (Jacq.) Ker Gawl.,

but it has come to our attention that /. purpurea Jacq. is

a later homonym for 7. purpurea Lam. (1789). Dating

of Jacquin ’s plates is complicated because the dates on

the title pages of the three volumes of the leones do not

represent the first publication of the individual plates,

which were released in fascicles earlier than the bound

volumes. The dating of the fascicles and the plates

included in each was resolved by Schubert (1945). Some

fascicles were predated by the publication of Jacquin ’s

Collectanea and the dates on the title pages of the sev-

eral volumes evidently do not reflect the actual year of

publication. Ixia purpurea Jacq. was first published in

Collectanea vol. 3, published in 1791 (title page 1789).

Fascicle 9 of the leones, which included 7 purpurea

Jacq. was, according to Schubert’s analysis, published in

1793.

Jacquin’s species is thus evidently predated by 7. pur-

purea Lam. (1789), a valid and legitimate name typified

by the plant in the Thunberg Herbarium named 7 crocata

var. c, flore purpurea. Lamarck was evidently providing

a name to one of the variants listed by Thunberg (1783)

in his account of Linnaeus’s 7 crocata (now Tritonia

crocata (L.) Ker Gawl.), thus excluding typical 7 cro-

cata (which Lamarck recognized as such), and 7 purpu-

TABLE 2. — Comparison of flowers of typical Babiana tubiflora and the Jacobsbaai population. Only well pressed, fully open flowers were

measured.

Bothalia 42,2 (2012)

143

rea Lam. is not nomenclaturally superfluous. Thunberg’s

I. crocata var. c was later regarded by De Vos (1983) as

merely a purple-flowered variant of Tritonia crocata.

The name Babiana purpurea is to be treated as a

new name (McNeill et al. 2006: Art. 58) dating from its

apparent transfer to Babiana by Ker Gawl., thus becom-

ing B. purpurea Ker Gawl. (1807a).

Babiana purpurea Ker Gawl. in Curtis’s Botanical

Magazine 26: sub. t. 1019 (1807a), nom. nov. pro Ixia

purpurea Jacq., Coll. 3: 268 (1791, as ‘1789’), horn, ille-

git. non Lam. (1789). Gladiolus purpureus (Ker Gawl.)

Willd.: 198 (1797). Gladiolus purpureus (Jacq.)Vahl:

114 (1805), horn, illegit. non Willd. (1797). Babiana

stricta var. purpurea (Ker Gawl.) Ker Gawl.: (1807b)

[name misapplied to B. villosa (Gmelin) Ker Gawl. ex

Steud.]. Type: South Alfica, without precise locality or

collector, illustration in Jacq., leones Plantarum Rari-

orum 2: t. 286 (1793).

3. Babiana bainesii Baker: in our revision of Babi-

ana (Goldblatt & Manning 2007a) we cited B. schlech-

teri Baker (1904) as a synonym of B. bainesii Baker

(1876). Unfortunately, we overlooked the fact that this is

a homonym for B. schlechteri Baker (1901) (= 5. nana

subsp. maculata (Klatt) Goldblatt & J.C.Manning). We

also neglected to note that the replacement name B. bak-

eri Schinz (1906) had been provided for Baker’s second

B. schlechteri. This correction has no nomenclatural

consequences at present. These names must be added to

the synonymy of B. bainesii as follows:

Babiana bainesii Baker in Journal of Botany 14: 335

(1876). Type: South Africa, [Gauteng,] ‘Gold Fields’,

Witwatersrand, near Johannesburg, 1870, Baines s.n. [K,

lecto.!, designated by Goldblatt & Manning: 92 (2004)].

Babiana bakeri Schinz: 712 (1906), as nom. nov.

pro B. schlechteri Baker: 1005 (1904), horn, illegit. non

Baker (1901). Type: South Africa, [Mpumalanga], near

Middleburg, 22 Dec. 1893, Schlechter 4055 (Z, ?holo.).

4. Babiana gariepensis Goldblatt & J.C.Manning:

currently known from three collections from the Rich-

tersveld in Northern Cape, B. gariepensis is a relatively

poorly known species of Babiana sect. Teretifolieae

(Goldblatt & Manning 2007a: 31). It is distinguished

by the unusually broad leaves mostly 70-110 x 18-25

mm, a stem produced shortly above the ground, pale,

grey-green flowers, the lower tepals with white streaks

edged dark red, and a relatively long perianth tube

20-24 mm long. A new collection of the species has now

come to our attention from southwestern Namibia in the

Namuskluft hiking trail near Rosh Pinah. Although a

modest range extension, some 50 km north of the near-

est station in South Africa, this represents the first record

of the species in Namibia. The one of the two specimens

of the collection is somewhat unusual for the species in

being particularly robust and has a stem reaching ± 120

mm above the ground; including the spike the specimen

is 280 mm tall.

Additional specimen

NAMIBIA. — 2716 (Witputz): Rosh Pinah, Namuskluft, on hik-

ing trail from campsite, (-DD), 27 July 2QW, R. & R. Saunders s.n.

(NBG).

5. Babiana lapeirousioides Goldblatt &

J.C.Manning: described in 2007 and based on a speci-

men and painting of a plant collected by C.L. Leipoldt

and grown to flowering at the National Botanical Gar-

dens, Kirstenbosch, in 1943, B. lapeirousioides was then

known only from the cryptic locality ‘Gariep’ (Goldblatt

& Manning 2007a: 49). This distinctive, long-tubed,

white flowered species was discovered in the wild by

Helga van der Merwe and Gretel van Rooyen in Octo-

ber 2011 at Vaalputs, the Nuclear Waste Disposal site

(between Gamoep and Platbakkies) in Northern Cape.

This collection allows us to expand the description of

the species, now with a specific locality (we associated

the original locality, Gariep, with an area of the Rich-

tersveld northeast of Port Nolloth and that now seems to

be mistaken and we wonder if ‘Gariep’ was a mistran-

scription of Gamoep).

Babiana lapeirousioides Goldblatt & J.C.Manning

in Strelitzia 18: 49 (2007). Type. — South Africa, [North-

ern Cape,] without precise locality, as ‘Gariep’, 30 Sept.

1943, Leipoldt s.n. (NBG, holo.).

Plants (5-) 10-20 cm high including leaves, stem

reaching shortly up to 70 mm above ground level, usu-

ally simple, rarely with 1 branch. Leaves sword-shaped,

blade 50-150 x 5-9 mm, rigid, plicate, pungent, smooth,

primary veins thickened and yellowish. Bracts (17-)20-

25(-35) mm long, glabrous or minutely scabrid above,

closely veined, dry and brownish above, inner slightly

shorter than outer, forked apically. Flowers in a com-

pact, 2^-flowered spike, zygomorphic, white with red

or purplish markings near base of lower tepals, presence

of scent unknown; perianth tube 22-3 5(— 40) mm long,

cylindric and straight, flaring slightly toward the mouth;

tepals subequal or upper three slightly larger, spreading,

narrowly oblanceolate, 10-20 x mm. Stamens uni-

lateral; filaments erect, 7-8 mm long; anthers 4. 5-5. 5

mm, cream-coloured. Ovary smooth; style dividing

from near base of anthers to shortly beyond anther tips,

branches ± 2.5 mm long. Flowering time: late Sept.-

early Oct.

Distribution and ecology: until now known only from

the type, the species has been recollected at Vaalputs

east of the Kamiesberg at the transition into Bushman-

land. The type locality, ‘Gariep’ is unlocalized and

we assumed it to refer to the Richtersveld but in the

light of the recent collection we consider that it might

equally refer to one or other of the inselbergs in south

of the Orange River in Bushmanland. The species grows

wedged in cracks among granite boulders, sometimes

forming large clumps. When not in flower the rigid, pun-

gent leaves are readily confused with B. dregei which

has a similar habitat in granite outcrops but at higher

elevations in the Kamiesberg.

Additional specimen

NORTHERN CAPE. — 3018 (Kamiesberg): Vaalputs, NECSA site,

(-AB), 1 Oct. 2011, G. van Rooyen 3048 (NBG, PRE).

6. Geissorhiza demissa Goldblatt & J.C.Manning:

most closely resembling G. aspera in its minutely

puberulous stems and radially symmetric flowers with

only slightly unequal filaments, G. demissa has rela-

tively small, white flowers tinged violet at the tips of the

144

Bothalia 42,2 (2012)

tepals (Goldblatt & Manning 2009). A new collection,

flowering after fire in the Kamiesberg, that we refer to

G. demissa, is unusually robust with larger flowers than

recorded until now for the species, the tepals ± 10 mm

long, anthers ± 3 mm long and up to four flowers per

spike. Floral dimensions previously recorded were tepals

7 X 3.5 mm and anthers ± 2 mm long and spikes had 2

or 3 flowers. The distinction between G. demissa and G.

aspera is less clear for this population. We revise a por-

tion of the key to subg. Geissorhiza (Goldblatt & Man-

ning 2009) below.

5a Stamens unequal, one filament at least 0.5 mm shorter than

other two:

6a Leaves ± plane with margins and central vein slightly to

moderately thickened, but not obviously winged; mar-

gins and central vein smooth or minutely puberulous:

7a Perianth predominantly purple, pale in throat; short fila-

ment at least 4 mm shorter than long filaments; tepals

18-23 mm long 81, G. inaequalis

7b Perianth predominantly blue to violet or white with pale

throat or purple with dark purple throat; short filament

no more than 2 mm shorter than long filaments; tepals

7-20 mm long:

8a Flowers radially symmetric except for eccentric style;

tepals 7-11 mm long; flowers violet to pale blue,

purple or predominantly white:

9a Tepals blue to violet with white throat edged in dark

violet, or uniformly white; mostly 11-14 x 4-6

mm; main spike mostly with 3-7 flowers; anthers

3-5 mm long 79. G. aspera

9b Tepals white, sometimes flushed dull purple outside

and tepals tips light violet, 7-10 x 3. 5,^.0 mm;

main spike with 1^ flowers; anthers 2-3 mm long

80. G. demissa

Additional specimen

NORTHERN CAPE. — 3018 (Kamiesberg): Langkloof, slopes

of Rooiberg between Farms Karas and Nartjiesdam, (-AD), 22 Sept.

2010, Goldblatt & Porter 13579 (MO, NBG)

ACKNOWLEDGEMENTS

Support for this study by grants 7103-01, 7316-02,

7799-05 and 8248-07 from the National Geographic

Society is gratefully acknowledged. We extend our grati-

tude to Elizabeth Parker and Lendon Porter for their

assistance and companionship in the field and Nick

Helme for bringing his discovery of the new Babiana

rivulicola to our attention. Collecting permits were pro-

vided by the Nature Conservation authorities of North-

ern Cape Province.

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Bothalia42,2: 147-186(2012)

A taxonomic revision of the southern African native and naturalized

species of Silene L. (Caryophyllaceae)

J.C. MANNING* and P. GOLDBLATT"

Keywords: Caryophyllaceae, new species, seed morphology, Silene L., southern Africa, taxonomy

ABSTRACT

The native and naturalized species of Silene L. in southern Africa are reviewed, with full synonomy and the description

of two new species from the West Coast of Western Cape. Eight native species and three naturalized species are recognized,

including the first identification in southern Africa of the Mediterranean S. nocturna L. The identity of 5. aethiopica Burm.,

which has remained unknown since its description, is established and is found to be the oldest name for S. clandestina Jacq.

Patterns of morphological variation within each species are discussed and subspecies are recognized for geographically seg-

regated groups of populations that are ± morphologically diagnosable. The following new names or combinations are made

among the southern African taxa: S. aethiopica subsp. longiflora; 5. burchellii subsp. modesta, subsp. multiflora, and subsp.

pilosellifolia; S. crassifolia subsp. primuliflora; S. saldanhensis; S. rigens; and S. undulata subsp. polyantha. Each taxon

is described, with information on ecology and distribution, and most species are illustrated, including SEM micrographs of

the seeds.

INTRODUCTION

Silene L. (tribe Sileneae), with 600-700 species

(Greuter 1995b), is distributed mainly through the tem-

perate regions of the northern Hemisphere, with its prin-

cipal centre of diversity in the Mediterranean and Mid-

dle East (Oxelman & Liden 1995). Over 90 species are

recorded in Africa, the vast majority in North Africa,

with very few extending southwards into sub-Saharan

Africa. Just three native and one introduced species of

Silene are known from south tropical Africa (Turrill

1956a; Wild 1961), with several additional species cur-

rently accepted in southern Africa.

Consensus has still to be reached on the delimita-

tion of Silene. The circumscription of the genus had

been gradually expanded over the last few decades of

the twentieth century until it coincided essentially with

the delimitation of tribe Sileneae (Greuter 1995b). This

consolidation was supported by a preliminary molecu-

lar sequence analysis of the tribe (Oxelman & Liden

1995), which nested all previously recognized genera

of Sileneae in Silene, with the possible exception of

Agrostemma L., but had been presaged by the break-

down of morphological differences between recognized

segregates when these were examined on a global scale.

Cucubalus L., which is deeply nested in Silene, is dis-

tinguished primarily by its berry-like fruits, which evi-

dently represent a specialized dispersal strategy. Lychnis

L., which forms a discrete clade within Sileneae, was

traditionally diagnosed by having capsules with as many

teeth as styles (vs. twice as many in Silene) (e.g. Chow-

dhuri (1957) and Chater & Walters (1964)) but the two

genera were diagnosed only with great difficulty by Bit-

■ Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, Claremont 7735, South Africa / Research Centre for

Plant Growth and Development, School of life Sciences, University of

KwaZulu-Natal, Pietermaritzburg, Private Bag XOl, Scottsville 3209,

South Africa. E-mail: J.Manning(gsanbi.org.za.

■■ B.A. Krukoff Curator of African Botany, Missouri Botanical Garden,

P.O. Box 299, St. Louis, Missouri 63166, USA. E-mail: peter.gold-

blatt(gmobot.org.

MS. received: 2011-12-19.

trich (1993) using a combination of trivial characters,

and his recognition of the two appears to have been pro-

visional until Silene could be conserved against Lychnis,

under which name it had earlier been included by Sco-

poli (1771). The generic name Lychnis therefore took

priority until it was formally rejected in favour of Silene

(Brummit 1994).

There are currently two opposing views on generic

delimitation in Sileneae. The first, advocated by Greuter

(1995b), is an inclusive one that recognizes just the two

genera Agrostemma and Silene. In this sense, Silene is

broadly distinguished within subfamily Caryophylloi-

deae (Bittrich 1993) by its actinomorphic flowers with

3 or 5 styles. The alternative view, recently developed

by Oxelman et al. (2000), proposes the partitioning

of Silene among seven genera, of which only Lychnis

(30 spp.) includes more than a handful of species. The

clade that constitutes Silene in this narrow sense is only

weakly supported in the molecular analysis and is also

not easily diagnosed morpholgically from its segregates.

Although we favour a more inclusive circumscription of

the genus, the native and introduced southern African

species are all included in Silene either way.

We follow the infrageneric classification of Silene

s. lat. proposed by Greuter (1995b), based largely on

Chowdhuri (1957) with some rationalization and sev-

eral nomenclatural corrections. Although there is mount-

ing evidence (Petri & Oxelman 2011) that at least some

of these sections are not monophyletic, no satisfactory

alternative has yet been offered. Significant characters

at sectional level are the type of inflorescence, presence

or absence of septa in the ovary, number of styles, shape

and venation of the calyx, number of teeth in the dehisc-

ing capsule, and the shape of the seeds.

Four global treatments have laid the foundations

for our present understanding of the genus (Otth 1824;

Rohrbach 1869, 1869-1870; Williams 1896; Chow-

dhuri 1957), and several modem regional treatments of

the European species have also been published (Greuter

1995b and references therein). Regional floristic studies

148

Bothalia42,2 (2012)

exist for parts of North Africa (Wickens 1976; Gilbert

2000) and much of sub-Saharan Africa (Turrill 1956a;

Wild 1961) but the taxonomy of Silene in southern

Africa still rests largely on Sonder’s (1860) treatment in

Flora capeusis, and only minor advances in our under-

standing of the southern African species have been made

since then (Rohrbach 1869, 1869-70; Bocquet 1977;

Masson 1989; Goldblatt & Manning 2000). The South

African species of Silene are unique among the flora

of the subregion in having been fundamentally misun-

derstood by Thunberg (1794), who misidentified the

majority of his collections as European species (Table

1 ), leaving it to later authors to recognize that they were

indigenous to the country and undescribed. Burman

(1768) had earlier made the same mistake, identifying as

European all but two {S. crassifolia L. and S. aethiopica

Burm.) of the eight species that he listed from the Cape.

Two previous attempts at revising the southern African

species, by G. Bocquet in the 1970s and D. Masson in

the 1980s, were never completed but many of the south-

ern African herbarium collections bear annotation labels

reflecting Masson’s unpublished species concepts and

names, adding further to the confusion. This revision of

Silene in southern Africa aims to redress the situation

and provide a base for future studies of the genus in the

subcontinent. This is especially important as the South

African winter rainfall region is a minor centre of diver-

sification for Silene in Africa.

Species delimitation in the genus is notoriously dif-

ficult and most African species, especially those with

a wider distribution, display a complex and often con-

fusing pattern of local variation that defies formal clas-

sification (Gilbert 2000). The primary reason for this is

almost certainly that the species are often facultatively

autogamous, which facilitates the differentiation of local

phenotypes, probably independently and repeatedly.

Population genetic studies are likely to shed significant

light on the nature and origin of these local forms. We

have recognized as species only entities that are mor-

phologically (and preferably also ecologically) discon-

tinuous, based on the assumption of ± complete inter-

specific infertility. Although Stace (1978) observed that

interspecific hybridization is generally rare in Silene,

recent molecular investigations (e.g. Erixon & Oxelman

(2008) and Frajman et al. (2009)) suggest that it is an

important consideration in some species groups. Where

possible, we recognize infraspecific taxa for geographi-

cally coherent groups of populations that show evident

but incomplete morphological differentiation. We have

uniformly applied the rank of subspecies to these taxa

but experimental investigation might indicate that other

ranks may be useful in some instances, such as in S.

aethiopica, where subsp. longiflora may be better treated

at the level of forma. The genetics of the maritime forms

of S. aethiopica and S. imdiilata also require investiga-

tion. Our intention, therefore, is to present a preliminary

classification that can form the basis for future experi-

mental studies in the genus.

Another intriguing aspect of the genus is its history

and diversification in southern Africa. The indigenous

species are currently distributed among three sections,

all of which are best represented in the northern hemi-

sphere and presumably originated there. Silene hurchel-

lii (sect. Fruticulosae) is widely distributed through the

eastern half of the continent, from Ethiopia (and Arabia)

to the Cape Peninsula in the extreme south, and the most

parsimonious interpretation is that the species migrated

from North Africa southwards along the East African

Afromontane corridor. Silene imdnlata (sect. Elisan-

the), in contrast, is largely southern African, extending

only as far north as the highlands of eastern Zimbabwe,

and its origin in southern Africa remains unclear but

long-distance dispersal appears to be more likely. Both

sections have subsequently undergone minor adaptive

radiation in the coastal regions of the subcontinent. The

third lineage is represented by S. aethiopica (sect. Dip-

terospermae), the only native southern African annual

species. Essentially endemic to the winter-rainfall region

in the extreme south and southwest, with no track in

tropical Africa, S. aethiopica most likely arrived in the

subcontinent through long-distance dispersal.

MATERIALS AND METHODS

All relevant types were examined, as well as all her-

barium material from BOL, MO, NBG, PRE, and SAM

(acronyms after Holmgren et al. 1990), the primary col-

lections of species from southern Africa. All species

were also studied in the field.

Ripe seeds from dehisced capsules were mounted

directly on aluminium stubs and coated with gold-pal-

ladium before viewing at the SEM Unit, University of

Cape Town. Seeds were sampled from at least two dif-

ferent collections per species (Figures 1, 2).

TAXONOMY

Silene L., Species plantarum: 416 (1753), nom. cons,

against Lychnis [Brummit: 272 (1994)]. Lectotype:

TABLE I — South African Silene species in the Thunberg Herbarium (UPS-THUNB). Native South African taxa in bold.

Bothalia 42,2 (2012)

149

FIGURE 1 . — Seed morphology and testa sculpturing in non-native southern Afncan Silene sect. Behen and sect. Silene. A, B, S. vulgaris, Moffet

613 (NBG): A, side; B, back. C, D, S. nocturna, Goldblatt & Porter 13551 (NBG): C, side; D, back. E, F, S. gallica, Bohnen 6351 (NBG):

E, side; E, back. Scale bar: 100 pm.

Silene anglica L., nom. rej. in favour of S. gallica L.,

designated by Britton & Brown; 62 (1913) = 5. gallica

L., nom. cons.

Cucubalus L.: 414 (1753). Type: Cucubaliis baccifer

L. = Silene baccifera (L.) Roth.

Lychnis L.: 436 (1753), nom. rej. in favour of Silene

L. [Brummit: 272 (1994)]. Lectotype: Lychnis chalce-

donica L., designated by Britton & Brown: 62 (1913) =

Silene chalcedonica (L.) E.H.L. Krause

Viscaria (DC.) Rohl.: 37 (1812). Lychnis sect. Vis-

caria DC.: 761 (1805). Type; Viscaria vulgaris Rohl. =

Silene viscaria (L.) Jess.

Annual, biennial or perennial herbs, rarely geophytes

or small shrubs, sometimes caespitose, rarely scandent,

glabrous or variously pubescent, sometimes glandular-

pubescent, rarely gynodioecious or dioecious. Leaves

opposite, sometimes slightly connate at base, sessile or

shortly petiolate, exstipulate. Inflorescence a dichasium

or monochasium, rarely 1 -flowered or sometimes ±

capitate. Flowers erect, spreading or pendulous, usually

bisexual, petals and stamens basally connate with floral

axis forming a short anthophore \fide Greuter 1995b].

Calyx tubular or dilated, sometimes strongly inflated,

5-toothed, 10(-60)-nerved, usually longitudinally ribbed

and ± plicate. Petals 5(0), usually white or pink, clawed,

limb entire to deeply lobed, mostly variously bifid, often

with paired coronal scales at base of limb, claw glabrous

or ciliate. Stamens 5 + 5 in male or bisexual flowers.

Ovary shortly stipitate, unilocular but usually 3(-5) sep-

tate in lower part, styles 3(6 or 10). Carpel subglobose

to ovoid, borne on persistent carpophore [anthophore],

cartilaginous and dehiscing apically by usually twice as

many (rarely as many) teeth as styles, rarely berry-like.

Seeds numerous, ± reniform, compressed, sometimes

disc-like and peripherally winged, variously scupltured,

often ± colliculate.

600-700 spp., mainly temperate Eurasia (± 600 spp.),

Africa (± 90 spp.) and N America (± 65 spp.).

Key to species [introduced taxa marked with an

asterisk]

la Inflorescence a ± divaricately branched dichasium; plants perennial:

2a Plants entirely glabrous, ± glaucous; calyx inflated, papery,

loosely investing capsule, equally 20-veined with veins

not raised; petal limbs divided to base, coronal scales

obsolete, claw abruptly narrowed and filiform in lower

half; ovary distinctly septate (sect. Behen) .... 1.5. vulgaris*

2b Plants pubescent with eglandular and glandular hairs;

calyx not inflated, unequally veined with median veins

thickened and prominent; petal limbs bifid, coronal

scales well-developed, claw gradually tapering to base;

ovary unilocular (sect. Elisanthe):

3a Lower portion of stem villous with spreading or shaggy,

eglandular hairs 1-2 mm long, sometimes mixed with

shorter eglandular or glandular hairs, upper part of

stem partly or entirely glandular-haired; calyx vestiture

partly or entirely eglandular (rarely entirely glandular),

lobes always fringed with eglandular hairs; stamens

and styles often ultimately well-exserted; carpophore

(2-)4— 10(-15) mm long; widespread through southern

Africa 2. 5. undulata

3b Stem either entirely glandular-haired or with a mix of

gland-tipped hairs and short, eglandular hairs 0.25-

0.50 mm long; calyx vestiture entirely glandular and

lobes fringed with glandular hairs; stamens and styles

included or very shortly exserted; carpophore 1-A mm

long; coastal sands and calcrete in extreme southwest-

ern Western Cape:

4a Stems stiffly erect, vestiture entirely of gland-tipped

hairs, without admixture of acute, eglandular hairs;

flowers relatively small, claws 8-10 mm long; cap-

150

Bothalia 42,2 (2012)

FIGURE 2. — Seed morphology and testa sculpturing in southern African Silene sect. Elisanihe. A, B, 5. uiidulata: A, side; B, back, Goldblatt 13549

(NBG). C, D, S. saldanhensis, Goldhlall & Manning 13646 (NBG): C, side; D, back. E, F, 5. ornata, Goldblatt & Porter 13254 (NBG): E,

side; F, back. G, H, S. rigens, Goldblatt & Porter 13291 (NBG): G, side; H, back. Scale bar: 100 pm.

Bothalia 42,2 (2012)

151

sules broadly um-shaped, 9-10 mm diam. at matu-

rity, causing calyx to split longitudinally at least mid-

way but often to base; seeds echinate 5. 5. rigens

4b Stems sprawling, vestiture a mix of gland-tipped hairs

and short, eglandular hairs; flowers larger, claws

15-18 mm long; capsules um-shaped, 6-8 mm

diam., not causing calyx to split; seeds colliculate or

tuberculate:

5a Plants mostly laxly sprawling, rarely compact in

exposed situations, with flowering stems strag-

gling; calyx without prominent, thickened ribs;

petals deep carmine, limbs not overlapping; seeds

tuberculate; growing on limestone outcrops in

rocky pavement 3. 5'. ornata

5b Plants compact, ± cushion-like with flowering stems

decumbent or erect; calyx with prominent, thick-

ened ribs; petals mauve, limbs overlapping; seeds

colliculate; growing in deep, calcareous sands and

consolidated dunes 4, 5. saldanhensis

lb Inflorescence a raceme-like monochasium, rarely only 1- or

2-flowered; plants annual or perennial:

6a Annuals; upper stems and inflorescence axes glandular-

pubescent; calyx um-shaped with triangular or awl-

shaped lobes, ribs bristly; carpophore < 1 mm long;

seeds compressed and peripherally winged with back

flat or shallowly grooved, testa colliculate or tuberculate

(sect. Silene):

7a Calyx glandular-pubescent and with glassy bristles 2-3

mm long on veins, without anastomising venation; pet-

als entire 1 1 . 5. gallica*

7b Calyx appressed-pubemlous and eglandular, with only

slightly longer hairs to 0.5 mm long on veins, anas-

tomising venation developed in distal half at least; pet-

als bilobed 10. 5. nocturna*

6b Perennials or annuals; stems and inflorescences eglan-

dular-pubemlous; calyx clavate with ovate lobes, uni-

formly shortly pubemlous; carpophore 2-12 mm long;

seeds compressed and disc-like, peripherally winged

with back deeply and acutely grooved between wings,

testa striate, cells radially fusiform:

8a Annuals (sect. Dipterospermae) 9. S. aethiopica

8b Geophytic perennials (sect. Fniticidosae):

9a Plants densely and closely branched, forming small

prostrate mats or loose tangled mounds with the

FIGURE 3. — Seed morphology and testa sculpturing in southern African Silene sect. Fruticulosae and sect. Dipterospermae. A-C, S. burchellii

subsp. burchellii, Penfold 111 (NBG): A, side; B, back; C, detail of dorsal groove. D-F, S. crassifolia subsp. crassifolia, Goldblatt et al.

13447 (NBG): D, side; E, back; F, detail of dorsal groove. G, S. crassifolia suhsp. primuliflora, Middelmost 2046 (NBG): G, side. H, I, S.

mundiana. Van der Merwe 1871 (NBG): H, side; I, back. J, K, 5. aethiopica subsp. aethiopica ‘typical form’, Goldblatt et al. 13469 (NBG):

J, side; K, back. L, M, S. aethiopica subsp. aethiopica ‘maritime form’, Goldblatt & Porter 13569 (NBG): L, side; M, back. Scale bar: A, B,

D, E, G-M, 100 pm; C, 10 pm; F, 30 pm.

152

Bothalia 42,2 (2012)

numerous inflorescences extending only shortly

above the vegetative growth; inflorescence axis

± filiform, 0.2-0. 5 mm diam.; leaves 7-15 x i_3

mm; racemes 1- or 2-flowered 8. .5. mimdiana

9b Plants not fomiing tangled mounds or mats and with

few inflorescences extending well above vegeta-

tive growth; inflorescence axis terete, 0.5-1. 5 mm

diam.; leaves 10-60 x (l-)2-12 mm; racemes

mostly more than 2-flowered:

I Oa Stems mostly erect or suberect, rarely decumbent,

mostly annual, 0.5-1 .5 mm diam. at base of inflo-

rescence; leaves linear to oblanceolate, rarely

obovate, thin-textured, clustered in lower half of

stem and well separated from inflorescence; calyx

weakly pleated; capsule ovoid, 8-10 x 5-7 rnm;

plants inland or coastal but never on foredunes

6. S. burchellii

1 Ob Stems ± prostrate or decumbent, relatively robust,

perennial and often branching, mostly 1. 5-2.0

mm diam at base of inflorescence; leaves oblan-

ceolate to suborbicular, ± succulent, running into

inflorescence; caljoc strongly pleated; capsule

broadly um-shaped to subglobose, (8-)10-12 x

7-10 mm; plants coastal on sandy foredunes ....

7. S. crassifolia

Silene sect. Behen Dumort., Florula belgica: 107

(1827). Sect. Inflatae (Boiss.) Chowdhurv. 241 (1957),

superfluous name. Silene [unranked] Inflatae Boiss.: 573

(1867). Type: Silene vulgaris (Moench) Garcke

± 8 spp., Europe and extratropical Asia, N Africa and

Atlantic islands; naturalized elsewhere.

Perennials or biennials, glabrous to sparsely eglan-

dular-pubescent, often glaucous. Leaves lanceolate to

orbicular. Flowers solitary or in few-flowered dicha-

sia, half-pendent at anthesis, vespertine (sometimes

indistinctly so); anthophore glabrous. Calyx glabrous,

inflated, 10- or 20-veined, with conspicuous reticulate

venation. Petals: claw glabrous, auriculate, limb bifid

to base; coronal scales linear or reduced. Stamens: fila-

ments glabrous. Ovary partially septate. Capsule subglo-

bose. Seeds semicircular-reniform, with flat to convex

flanks and back, radially colliculate-echinate.

1. Silene vulgaris (Moench) Garcke, Flora von Nord-

and Mittel-Deutschland, ed. 9: 64 (1869). Cucubalus

behen L.: 414 (1753), non Silene behen L. (1753). Behen

vulgaris Moench: 709 (1794). Silene cucubalus Wibel:

241 (1799), nom. illegit. superfl. Silene inflata Sm.: 467

(1800), nom. illegit. superfl. Type: ‘Habitat in Europae

septentrionalis pratis siccis’. Herb. Linn. 582.4 [LINN,

lecto.l, designated by Aeschimann & Bocquet: 204

(1983)].

[see Chater & Walter (1964) and Greater (1997) for

heterotypic synonyms]

Gynodioecious perennial to 60 cm tall, softly woody

at base, stoloniferous; stems ascending or erect, 1-3

mm diam., usually branched, glabrous, with or without

axillary non-flowering shoots. Leaves ovate-lanceolate

or the lower oblanceolate, 30-90 x 6-15 mm, acute to

apiculate, base cuneate or narrowed and petiole-like

in lower leaves, glabrous, glaucous, with evident side

veins. Inflorescence a laxly divaricate dichasium, usu-

ally several-flowered; bracts smaller than leaves, ovate-

attenuate, up to 1 5 X 5 rnm, thinly herbaceous with sub-

membranous margins or entirely ± scarious; pedicels

mostly 10 15 mm long at anthesis but up to 30(-50) mm

long in fruit. Calyx inflated, um-shaped and bladder-like,

10-12 X 7-8 mm at anthesis but accrescent to 13x 10

mm in fruit and loosely investing capsule, base intm-

sive, whitish or pale green with darker veining, equally

20-veined with reticulate venation throughout, not pli-

cate, lobes broadly triangular, 1.5 x 2.0 mm, densely

ciliolate along margins and inner faee toward apex.

Flowers spreading or ± pendent at anthesis, bisexual

or functionally female, lily-scented at night only but

remaining open during day. Petals white or pink to pur-

ple, claw 10-11 mm long, cuneate and conspicuously

auriculate in upper half, abmptly narrowed and filiform

in basal half, limb deeply bilobed almost to base, 4-6 x

4-5 mm, coronal scales obsolete. Stamen filaments ± 1 8

mm long and exserted ± 7 mm, or in functionally female

flowers 5-6 mm long and deeply included, glabrous.

Ovary ovoid, ± 3 mm long; styles 1 8 mm long, exserted

7-8 mm. Capsule um-shaped, ±10x7 mm, usually ± 3

X longer than carpophore; carpophore 2. 5-3.0 mm long,

glabrous. Seeds reniform with flat or convex face and

convex back, 1.0-1. 5 mm, tuberculate, grey. Flowering

time: Oct. -Dec. (-Jan.). Figures lA, B, 4.

Vernacular name: bladder campion.

Distribution and ecology: native to Europe, where

it is widely distributed across the continent through the

Middle East into temperate Asia; introduced into Ethio-

pia, Australia, South Africa (where it is cultivated as an

ornamental) and also North America, where it is consid-

ered a weed.

Diagnosis: readily distinguished by the complete lack

of vestiture and by the pale greenish or whitish, inflated,

bladder-like calyx with 20 equal longitudinal veins

joined by extensive reticulated veining throughout. The

petals, deeply divided to the base of the limb with the

claw abmptly narrrowed and filiform in the basal half,

are also diagnostic. Flowers are either bisexual, with

long, well-exserted filaments ± 18 mm long, or func-

tionally female, with relatively short filaments, 5-6 mm

long, deeply included within the calyx.

The species is broadly circumscribed by Chater &

Walters (1964), who recognize eight subspecies for the

European material, most of which have been variously

treated by other authors. The South African material is

stoloniferous and is thus treated as subsp. macrocarpa

Turrill (1956b).

subsp. macrocarpa Turrill, Hooker’s leones

plantarum 36: t. 3551 (1956b). Type: Cyprus, Chion-

istra, 1 620 m, 4 Jun. 1940 [grown in the Herbarium

Experimental Ground, Kew, from seeds collected 5 Jul.

1938], Kennedy s.n. K2312 (K, holo.).

Distribution and ecology: widely distributed through

the Mediterranean and long established in southwest

Britain, whence it was presumably introduced to South

Africa, probably as an ornamental. In South Africa the

species has been recorded from mesic situations in the

Western Cape winter-rainfall region (Tygerberg Hills,

around Stellenbosch and Oudtshoom, and in the Lang-

klool) and from the Hogsback in Eastern Cape, as a

garden escape in waste places, along roadsides, around

old lands, and in vineyards. It was first recorded as an

Bothalia 42,2 (2012)

153

FIGURE 4. — Silene vulgaris, ex

hort., no voucher. A, flower-

ing stem; B, petal; C, gyn-

oecium; D, ovary 1/s; E, seed.

Scale bar: A, 10 mm; B, C, 2.5

mm; D, 2 mm; E, 0.5. Artist:

John Manning.

escape around Stellenbosch in the early twentienth cen-

tury (Figure 5).

In South Africa, Silene vulgaris subsp. macrocarpa

can form dense stands, propagating through stolons but

always in transformed habitats and there is no indica-

tion that it is invasive. The extensive, stoloniferous root-

stock may however become a problem for cultivation

if the plants are allowed to establish unchecked over

several years in fallow lands, and subsp. macrocarpa

has become a weed of agronomic importance in Europe

(Aeschimann 1983).

Additional specimens

WESTERN CAPE. — 3318 (Cape Town): Tygerberg Nature

Reserve, (-DC), 12 Dec. 1975, Loubser 3476 (NBG); Rustenberg

154

Bothalia 42,2 (2012)

FIGURE 5. — Recorded distribution of Silene vulgaris in southern

Africa.

Farm, (-DD), 7 Oct. 1973, Taylor 8608 (NBG, PRE); Idas Valley,

Schoongesicht Farm, (-DD), 26 Oct. 1973, Neser s.n. (NBG); Stel-

lenbosch, Blaauwklip, (-DD), 10 Oct. 1928, Gillett 582 (NBG);

Stellenbosch, Devon Valley, (-DD), 21 October 1968, Du Toil s.n.

(NBG); Jonkershoek, (-DD), 19 Oct. 1943, Adamson 3514 (BOL).

3322 (Oudtshoom): Prince Albert, (-AA), 8 Nov. 1988, Dean 748

(PRE); Oudtshoom Experimental Farm, (—AC), 29 Jan. 1970, Maree

40 (NBG); Boomplaas, Cango Valley, (-AC), 5 Feb. 1975, Mojfetl

613 (NBG). 3323 (Willowmore): Longkloof, (-BC), 13 November

1970, Botha s.n. (NBG, PRE); Avontuur, (-CA), 2 Jan. 1970, Wild s.n.

(NBG)

EASTERN CAPE. — 3226 (Fort Beaufort): Hogsback, Risingham,

old cultivated land, (— DB), 10 Jan. 1989, King s.n. PRE60819 (PRE).

Silene sect. Elisanthe (Fenzl. ex Endl) Ledeb., FI.

Ross. 1: 314 (1842). Saponaria sect. Elisanthe Fenzl. ex

Endl., Gen. PL: 972 (1840). Silene sect. Melandriformes

(Boiss.) Chowdhuri: 244 (1957), superflous name,

Silene [unranked] Melandriformes Boiss., FI. Or. 1: 568

(1867). Lectotype, designated by Pfeiffer: 1186 (1871-

1875): Silene noctiflora L.

Annuals, biennials and perennials, pubescent with

eglandular and glandular hairs, monoecious or dioe-

cious. Leaves ovate-lanceolate; bracts herbaceous. Flow-

ers in few-flowered dichasia, bisexual or unisexual,

spreading to ± erect, mostly vespertine; anthophore gla-

brous or pubescent. Calyx pubescent, clavate, 1 0-veined,

with conspicuous reticulate venation. Petals', claw gla-

brous to ciliate, auriculate, limb bifid; coronal scales

present. Stamens', filaments glabrous or pubescent.

Ovary unilocular at maturity; styles 3 or 5. Seeds semi-

circular-reniform, with convex flanks and back, radially

colliculate-echinate.

± 20 spp., Eurasia and N Africa, tropical and S Africa.

The large-flowered southern African species were not

treated by Chowdhuri (1957) in an inexplicable omis-

sion, but have traditionally been placed in sect. Elisan-

the (Sonder 1860; Chater & Watters 1964), with which

they accord in their dichasial inflorescence, reticulate

calyx venation, completely unilocular ovaries, and col-

liculate-echinate seeds without peripheral wings.

2. Silene undulata Aiton, Hortus kewensis 2: 96

(1789); Sond.: 126 (1860). Silene tristis Salisb.: 301

(1796), nom. superfl. illegit. Melandrium ornatum var.

undulatum (Aiton) Rohrb.: 234 (1869). Melandrium

undulatum (Aiton) Rohrb.: 245 (1869-70). Type: South

Africa, without precise locality, cultivated at Kew

from seed introduced by Francis Masson, Masson s.n.

BM000593534 (BM, holo.!)

Tufted perennial, 0. 5-2.0 m, producing new vegeta-

tive shoots from basal axils at end of flowering season,

often developing a gnarled, softly woody crown; root-

stock a carrot-like taproot. Stems erect to sprawling,

rarely prostrate in exposed situations, simple to highly

branched, 1.5-5.0(-8.0) mm diam., sparsely or densely

pubescent with lower portions of stems bearing acute,

eglandular hairs sometimes mixed with scattered, short,

gland-tipped hairs, eglandular hairs either all longer or

a mix of longer, spreading or shaggy hairs 1-2 mm long

and shorter, decurved hairs, upper parts of stems usually

with progressively more gland-tipped hairs, sometimes

almost entirely glandular-haired. Leaves mostly basal,

suberect or spreading, lower leaves oblanceolate-spathu-

late, sub-petiolate, up to 200 x 25 mm, sometimes dead

or dying at flowering, cauline leaves elliptical to oblan-

ceolate, mostly 50-80 x 10-20 mm, acute or acuminate,

base tapering, margins plane or undulate, usually pubes-

cent, either with mixture of erect, gland-tipped hairs and

acute, eglandular hairs or entirely eglandular, sometimes

subglabrous or rarely glabrous except on margins, mar-

gins always ciliate, with longer hairs to 1 mm long at

extreme base, with evident side veins. Inflorescence a

lax or more compact, symmetrically or asymmetrically

branched cyme, axis 1-2 mm diam. below primary

flower; bracts similar to upper leaves but smaller, sub-

equal, suberect or spreading; primary pedicels 8^0 mm

long, secondary pedicels 5-20 mm long. Flowers half-

nodding, nocturnal, clove- or soapy-scented at night.

Calyx cylindrical or narrowly um-shaped in flower,

(18-)20-35(^5) mm long, 10-ribbed, with 1^ reticu-

late veins in distal half, densely or sparsely puberulous,

hairs usually a mix of acute, eglandular and spreading,

gland-tipped hairs but sometimes entirely glandular or

entirely eglandular, lobes narrowly triangular to awl-

shaped, (3-)5-10 mm long, densely ciliolate. Petals

white to pink with yellowish reverse, limbs spreading,

cuneate, not overlapping with adjacent petals, 9-15 x

8-15 mm, bifid ± halfway, claws strap-shaped, (12-)15-

25 X 2. 5-3.0 mm, glabrous or pubescent along abax-

ial midline and margins, exserted up to 10 mm beyond

calyx, auriculate apically, auricles of adjacent tepals

locking together, coronal scales spreading or suberect,

0.7-1. 5 mm long, denticulate. Stamen filaments unequal,

shorter series 11-25 mm long, longer series 16-30 mm

long, reaching to top of claws or exserted; anthers ± 2

mm long, reaching top of claws and included or exserted

up to 6 mm. Ovary narrowly pyriform, shortly stipitate,

5-7 mm long, stipe ± 1 mm long; styles ±9-15 mm long

at anthesis and included but usually elongating, ulti-

mately reaching up to 25 mm long and exserted up to

10 mm. Capsule ovoid, (10-) 15-25 x 5-8 mm, 1. 5-6.0

X longer than carpophore, minutely granular; carpophore

(2-)4-10(-15) mm long, glabrescent. Seeds 1.2-1. 5 mm

diam., reniform with hilum recessed, face flat, back flat,

reddish brown, testal cells radial-concentric, colliculate-

tuberculate. Flowering time'. Aug.-Dee. in the winter

rainfall region; Nov.-Mar. in the summer rainfall region

Bothalia 42,2 (2012)

155

but as late as June along the subtropical coast. Figures

2A, B, 6, 7.

Vernacular names: Cape catchfly, wild tobacco,

wildetabak.

Distribution and ecology: widely distributed through

the more temperate parts of southwestern and eastern

southern Africa northwards into eastern Zimbabwe (Fig-

ure 8), occurring in a variety of stony or rocky habitats,

in coastal and inland scrub, often in ravines, along river-

banks and forest margins, and in moist grassland, from

just above the high tide mark to well over 2 000 m. The

species is absent from the more arid regions, notably

Bushmanland and the Kalahari in Northern Cape and the

Great Karoo in Western and Eastern Cape.

Growth and flowering of the species in vegetated

communities such as thicket margins is stimulated by

clearance of the overburden by fire. The flowers are

strictly nocturnal and strongly scented at night but

although they may remain open for several hours on

overcast mornings, sometimes until midday, they are

then often no longer scented. The species is clearly pal-

atable, being extensively browsed by both wild animals

and livestock, but is also used traditionally as a medicine

in many diseases, particularly fevers and delirium (Watt

& Breyer-Brandwijk 1932).

Diagnosis and relationships: as circumscribed here,

Silene undulata is a highly variable species with numer-

ous local forms that have in common the development

of relatively long, eglandular hairs, 1-2 mm long, on the

lower portion of the stem, giving the base of the stems a

characteristic, shaggy appearance (Figure 7C). In some

instances the eglandular hairs are mixed with scattered,

short, gland-tipped hairs, which typically predominate

in the upper parts of the stems (Figure 7B), which are

sometimes entirely glandular-haired. The vestiture of

the calyx is similarly variable, usually including a mix

of longer, eglandular hairs and short, gland-tipped hairs,

but sometimes either entirely eglandular or entirely glan-

dular. The lobes are, however, invariably fringed with

eglandular hairs. Other sub-Saharan species in the sec-

tion have a ± entirely glandular pubescent calyx, includ-

ing the margins of the lobes. Another characteristic of

the species is the relatively long, narrowly triangular to

FIGURE 6. — Silene undulata subsp.

undulata, Clanwilliam, Gold-

blatt & Porter 13523. A,

flowering stem and flower;

B, calyx laid out; C, petal;

D, gynoecium before style

elongation, 1/s and t/s; E, gyn-

oecium after style elongation;

F, capsule after dehiscence; G,

seed. Scale bar: A, F, 10 mm;

B, 6 mm; C-E, 2.5 mm; G,

0.5. Artist: John Manning.

156

Bothalia 42,2 (2012)

FIGURE 7. — Silene unchilata subsp.

undidata, Langebaan, without

voucher. A, flowering stem; B,

peduncle segment; C, lower

stem segment; D, calyces laid

out; E, petal; F, gynoecium

before style elongation; G,

capsules after dehiscence (left

hand illustration with calyx

removed). Scale bar: A, G, 10

mm; B, C, 1 mm; D, 5 mm;

E, 2.5 mm. Artist: John Man-

ning.

subulate calyx lobes, mostly 5-8 mm long (Figures 6B,

7D).

The species varies also in the length and exsertion of

the stamens and styles. Style length is dependent on the

age of the flowers, and the styles are always included at

anthesis. In many populations, however, the styles elon-

gate markedly after anthesis, becoming well exsertcd by

the second or third day.

Silene unchilata was described from cultivated plants

grown at Kew from seeds collected in the Western Cape

by Francis Masson, who visited the region from 1772 to

1775 and again from 1786 to 1795. The earliest collec-

tion of the species, however, appears to be an unlocal-

ised specimen at the British Museum collected in 1772

by Franz Oldenburg {Oldenburg 1260). Oldenburg,

who visited the Cape inl771-1773, accompanied both

Masson and Carl Thunberg on collecting trips (Glen &

Germishuizen 2010). His fragmentary specimen at the

British Museum (BM001010634) was initially ignored,

then mistakenly identified as the Sicilian species Silene

calycina C.Presl., before finally being correctly deter-

mined as S. undulata. It represents the form with a pro-

portionally longer carpophore that was initially recog-

nized at species level under the name S. capensis Otth.

An equally fragmentary collection of the same form

made by Thunberg, possibly from the same population

that yielded seeds to Masson since the two men certainly

collected together on the Cape Peninsula and along the

West Coast where S. undidata occurs, was mistaken by

him for the Mediterranean S. pendula (Table 1 ).

Bothalia42,2 (2012)

157

FIGURE 8. — Distribution of Silene widiilata in southern Africa: subsp.

undulata, •; subsp. polyantha, o.

Silene imdiilata exhibits numerous local forms, vary-

ing in habit, degree and nature of vestiture, length of

calyx and carpophore, length and shape of calyx lobes,

size of petals, degree of exsertion of stamens and styles,

and shape and size of the capsule. Some of these have

formed the basis of separate species but we are unable

to identify any of these character states or combination

of states that is consistent with the recognition of more

than a single, variable species. The relative length of

carpophore and capsule, in particular, has been used to

distinguish several species in the complex, viz. S. bel-

lidioides (carpophore one quarter as long as capsule), S.

undulata (carpophore one third as long as capsule), and

S. capensis (carpophore at least half as long as capsule)

(Sonder 1860). A further two species were recognized on

the basis of their relatively hairless leaves, 5. eckloniana

from Cape Recife with eglandular stems, and leaves gla-

brous above but hairy on the margins and along the mid-

rib beneath, and S. cajfra, also from Eastern Cape, with

subglabrous stems and leaves that are ciliolate along the

margins but otherwise glabrous on both surfaces. With

numerous collections now to hand, we are unable to

correlate any of these differences with other character

states, nor with geography, and conclude that they repre-

sent independent and local variants within a single spe-

cies.

Some of these names were consolidated by Rohr-

bach (1869), who treated S. cajfra, S. capensis, and S.

diurniflora as synonyms of 5. undulata. Of the remain-

ing names, only S. bellidioides and S. undulata have

been applied in recent years, always with a great deal of

uncertainty reflecting the ± continuous and often local

variation in the relative lengths of carpophore and cap-

sule.

There is, however, a distinctive morphotype char-

acteristic of the populations from the coastal and near-

inland parts of KwaZulu-Natal and Swaziland. Plants

from this region consistently produce stiffly erect stems,

with closely branched, flat-topped cymes of flowers with

short calyces, mostly 12-15 mm long, and unusually

small capsules, 10-15 mm long. Although all of these

states are found elsewhere in the species, their consistent

association among these populations is unique and sug-

gests some degree of genetic differentiation. We recog-

nize these plants as subsp. polyantha.

Key to subspecies

la Stems erect or sprawling; cymes mostly without tertiary

branching; calyx ( 15-)20-35(^5) mm long; capsules

(1 2-) 15-25 mm long, 1.5^.0 x longer than carpophore;

plants widespread but not from coastal KwaZulu-Natal or

Swaziland 2a. subsp. undulata

lb Stems erect; cymes with tertiary and often quartemary

branching; calyx 12-15(-20) mm long; capsules 10-15

mm long, 4-6 x longer than carpophore; plants from

coastal KwaZulu-Natal or Swaziland 2b. subsp. polyantha

2a. subsp. undulata

Silene capensis Otth in DC.: 379 (1824); Sond.: 125

(1860). Melandrium capense (Otth.) Rohrb.: 232 (1869).

Melandrium undulatum var. capense (Otth.) Rohrb.: 246

(1869-70). Type: South Africa, ‘ad cap. B.-Spei.’, with-

out date or collector. Prodr. I p. 379 no. 144 (G-DC,

holo.-microfiche! ).

Silene diurniflora Kunze: 578 (1844) [fide Sond.

(I860)]. Type: South Africa, ‘Prom. b. sp.’, cultivated

at University of Leipzig 1842-1843, Guenzius s.n. (LZ,

holo.t).

Silene bellidioides Sond.: 125 (1860), syn. nov.

Melandrium bellidioides (Sond.) Rohrb.: 247 (1869-70)

[as "bellidioide']. Type: South Africa, ‘fields near the

Zwartkops River, Nov. without year, Ecklon & Zeyher

1964 S-G-5677 (S, lecto.!, designated by Nordenstam:

279(1980); SAM!, isolecto.].

Silene eckloniana Sond.: 126 (1860), syn. nov.

Melandrium ecklonianum (Sond.) Rohrb.: 232 (1869).

Type: South Africa, [Eastern Cape], ‘sea shore near

Cape Recife, Feb. 1830/2’, Ecklon s.n. S-G-8711 (S,

holo.i).

Silene caffra Fenzl ex C.Muell. in Walp.: 216 (1868).

Melandrium cajfrum (Fenzl ex C.Muell.) Rohrb.: 232

(1869). Melandrium undulatum var. cajfrum (Fenzl ex

C.Muell.) Rohrb.: 246 (1869-70). Type: South Africa,

[Eastern Cape], ‘Caffraria’, Drege 5342 (W, holo.t).

Silene bellidifolia var. stricta Fenzl in Drege: 222

(1844), nom. nud.

Silene bellidifolia var. foliosa Fenzl in Drege: 222

( 1 844), nom. nud. \fide Sond. ( 1 860)].

Silene caffra Fenzl in Drege: 222 (1844), nom. nud.

Silene capensis var. flaccida E.Mey. in Drege: 222

(1844), nom. nud.

Silene meyeri Fenzl in Drege: 222 (1844), nom. nud.

[et herb. Drege 730, fide Rohrb.: 245 (1869-70)].

Silene ornata lus. flaccida Fenzl in Drege: 222

(1844), nom. nud. [fide Sond. (1860), as fiorida'].

Silene thunbergii E.Mey. in Drege: 222 (1844), nom.

nud., non S. thunbergiana Eckl. & Zeyh. ex Sond.

(1860) [fide Sond. (I860)].

158

Bothalia 42,2 (2012)

Melandrium capense var. strictum [as "stricta''\ Fenzl

ex Rohrb.: 232 (1869), nom. nud.

Silene noctiflora sensu Thunb., in part [excluding

UPS-THUNB 10774 & 10775 = S. hgens J.C.Manning

& Goldblatt]: 81 (1794), non. L. (1753).

Stems erect or sprawling. Cymes relatively few-flow-

ered, mostly without quartemary branching, rounded.

Calyx (15-)20-35(^5) mm long. Capsules ovoid to

narrowly ovoid, (12-)15-25 mm long, l.S^.O x longer

than carpophore. Carpophore 4-10(-15) mm long. Fig-

ures 6, 7.

Distribution and ecology, throughout most of the

range of the species but replaced along the eastern sea-

board by subsp. polyantha (Figure 8).

Diagnosis and relationships', a variable taxon, typi-

cally with a lax, rounded inflorescence of relatively large

flowers with calyx mostly 20-35 mm long, often with

subulate lobes. The nature of the stem and calyx vesiture

is very variable, with some plants entirely eglandular and

others predominantly glandular, sometimes even within

a single collection (e.g. Taylor 5180). Variation in the

nature and degree of vesiture is especially evident among

the populations along the southern Cape coast. Popula-

tions between Knysna and Plettenberg Bay often, but not

always, have entirely eglandular stems and calyces.

Most populations have the anthers exserted beyond

the mouth of the floral tube at anthesis, followed by the

styles, which at anthesis are always included in the floral

tube. This feature is thus evident only in older flowers.

Also diagnostic of many populations are the long, subu-

late calyx lobes, 5-7 mm long but this can vary within

populations. Plants with short calyces, ± 15 mm long,

and carpophores ± 4 mm long, are typical of the West

Coast, whereas specimens from western Lesotho (e.g.

Schmitz 8095, Hilliard & Burtt 12065) have among the

longest calyces (35^5 mm long) and carpophores (± 15

mm long) recorded for the species.

Plants are very variable in habit, those from coastal

and alpine habitats typically more compact and pros-

trate whereas individuals from sheltered or shaded situ-

ations are more lax. Some of this variation is certainly

ecological and linked to the habitat but some may well

be genetic, since plants from the cliffs at Hermanns evi-

dently retain their compact habit in cultivation [Davis

s.n. (BOL)].

Separation from subsp. polyantha is not always clear

in the Kwazulu-Natal Midlands, especially around Pie-

termaritzburg, where plants have erect stems and well-

branched, flat-topped inflorescences but calyces ± 20

mm long.

Additional specimens

LIMPOPO.- 2330 (Tzaneen): Woodbush, (-CC), Jan. 1923,

Wager s.n. TRV23094 (PRE), 2429 (Zebediela): Donkerkloof near

Chuniespoort, I 700 m, (-BA), 14 Mar. 1974, Vahrmeijer 2414 (MO,

PRE).

NORTH-WEST.— 2527 (Rustenburg); Rustenburg, (-CC), 5 Nov.

1940, Lanham s.n. (PRE). 2624 (Vryburg): Vryburg, Aarbosvlaktc, (-

DC), 25 Mar. 1921, Mogg 8435 (PRE); Vryburg, (-DD), Apr. 1912,

Rogers s.n. HOL45405 (BOL).

GAUTENG. — 2527 (Rustenburg): Witwatersrand, Hekpoort, (-

DC), 18 Apr. 1936, Phillips 33 (PRE). 2528 (Pretoria): Pretoria, Arca-

dia, E of Union Buildings, (-AC), 12 Jul. 1955, Smith 323 (PRE);

Pretoria, (-CA), 21 Nov. 1904, Leendertz 4148 (PRE); Waterkloof,

(-CA), 25 Oct. 1920, Verdoorn 131 (PRE); Wonderboom Poort, (-

CA), 24 Nov. 1917, Pole-Evans 230 (PRE); Fountains Valley, (-CA),

14 Mar. 1930, Verdoorn 820 (PRE); Moreletta spruit, (-CA), 1 Mar.

1980, Germishuizen 1260 (PRE); Irene, (-CC), Apr. 1924, Smuts 1318

(PRE); Doomkloof, (-CC), 8 Nov. 1928, Gillett s.n. (NBG). 2627

(Potchefstroom): Klifdrif, E of Potchefstroom, (-AD), 8 Jan. 1935,

Theron 1208 (PRE); Parys, (-CC), Apr. 1907, Grey College Herbar-

ium 576 (BOL).

MPUMALANGA. — 2530 (Lydenburg): Verlorenvallei Nature

Reserve, NE of Dullstroom, (-AC), 16 Dec. 1932, Galpin s.n.

BOL45397 (BOL); 10 Apr. 2000, Van Slageren & Van Wyk 971 (PRE);

Mokobulaan Plantation on Skurweberg, 2 100 m, (-BA), 3 Dec. 1985,

Kluge 2663 (PRE). 2629 (Bethal): Ermelo, Nooitgedacht, (-DB), 5

Feb. 1927, Henrici 1467 (PRE). 2630 (Carolina): Carolina, (-AA),

Dec. 1905, Bolus 11701 (BOL).

FREE STATE. — 2827 (Senekal): Gumtree, (-DD), 13 Jan. 1948,

Reardon 4 (NU). 2828 (Bethlehem): Golden Gate Highland Park, (-

DA), Jan. 1966, Liebenberg 7495B (PRE); Qwa Qwa National Park,

Qwa Qwaberg, (-DB), 2 Feb. 1995, Zietsman 3112 (PRE). 2829 (Har-

rismith): Harrismith, Drakensberg Botanical Garden, (-AC), 16 Jan.

1974, Jacobsz 2015 (MO, NBG, PRE, S); foot of Platberg, (-AC), 2

Feb. 1968, Van der Zeyde 227 (NBG). 2926 (Bloemfontein): Bloem-

fontein, Gen. de Wet School, (-AA), 3 Jun. 1967, Hanekom 910

(PRE); O.F.S. Botanic Garden, (-AA), 27 Mar. 1968, Muller 236

(NBG); Mazelspoort, (-AA), 7 May 1949, Steyn 50 (NBG); Dewets-

dorp, (-DA), 15 Apr. 1950, Steyn 942 (NBG).’ 2927 (Maseru): Lady-

brand, (-AB), Nov. 1906, Rogers 5241 (PRE). 3025 (Colesberg): 5 km

from Colesberg on road to Steynsburg, (-CA), 1 Apr. 1981, Herman

465 (PRE). 3027 (Lady Grey): Zastron campsite, (-AC), 27 Mar. 1980,

Reid 202 (PRE).

KWAZULU-NATAL.— 2730 (Vryheid): Utrecht Dist., Groenvlei,

Balele Mountain Lodge, (-CA), 18 Nov. 1997, Ngwenva 1617 (NH);

Hlobane, (-DB), 5 Nov. 1950, Johnstone 5/7 (MO, NU). 2828 (Beth-

lehem): Witzieshoek, (-DB), Dec. 1905, Thode 5664 (NBG); Royal

Natal National Park, near Basuto Gate, (-DB), 2 Feb. 1982, Stewart

& Manning 2254 (NU). 2829 (Harrismith): Van Reenen, (-AD), Jun.

1914, Bews 965 (19U). 2929 (Underberg): Cathedral Peak Forest

reserve, (-AB), 1 Jan. 1983, Noel 2820 (NU); Giant’s Castle Game

Reserve, near Dinosaur footprints, (-AD), 27 Dec. 1967, Trauseld

907 (NU); Lotheni Valley, vicinity of Ash Cave, (-AD), 7 Feb. 1985,

Hilliard c6 Burtt 18188 (E, NU, PRE); Spring Grove Farm, (-BB/

BD), 9 Dec. 1996, Greene 906 (NH); Kamberg, (-BC), 28 Nov.

1974, Wright 1985 (NU); 10 Jan. 1990, Williams 687 (NH); Mpendle

Dist., Mulangane Ridge, (-BC), 3 Feb. 1984, Hillard & Burtt 1750 7

(E, K, NU, PRE, S); Chameleon Cave area, (-CB), 1 Dec. 1984,

Hilliard & Burtt 17769 (E, K, NU, PRE, S); Sani Pass, (-CB), 18

Feb. 1973, Hilliard 5339 (NU); 23 Mar. 1977, Hilliard & Burtt 9801

(E, K, MO, NU); 6 Jan. 1984, Hilliard, Burtt & Manning 17284 (E,

NU); Jan. 2000, Edwards 1749 (E, K, NU); Cobham State Forest,

Polela River, (-CB), 2 Dec. 1987, MacDevette 2036 (NH); Vergelegen

Nature Reserve, (-CB), 2 Jan. 1978, Hilliard Burtt 11182 (E, MO,

NH); Underberg, Watermead farm, (-DC), 16 Dec. 1989, Williams

645 (NH); Bulwer, Sunset farm, (-DC), 24 Feb. 1990, Vos 31 (NU);

Polela Dist., Hlabeni Mtn near Creighton, (-DC/DD), 21 Nov. 1994,

Wirminghaus 1310 (NH). 2930 (Pietermaritzburg): Mount West Dist.,

3 km N of intersection, (-AA/AC), 13 Nov. 1988, Greene 609 (NH);

Kunhardt’s Farm, 21 km from Merrivale on Boston Road, (-AC),

Jan.-Apr. 1982, Kunhardt 10 (NH); Karkloof Mtn Range, summit of

Mt Gilboa, (-AD), 18 Nov. 2000, Johnson & Neal 29 (NU); Pieter-

maritzburg, World’s View, (-CB), 30 Oct. 1964, Tunnington s.n. (NU);

Richmond/Ixopo area, Hella Hella, (-CC), 1 Nov. 1997, Swanepoel &

Porter 14 (NH). 3029 (Kokstad): Mzimkhulu Dist., Nsikeni Nature

reserve, (-AB), 19 Feb. 1992, Williams 901 (NH); Newmarket, (-AD),

1 Feb. 1895, Krooks.n. SlO-9935 (S); Mt Currie, (-AD), 20 Jan. 1957,

Taylor 5483 (NBG); between Flagstaff and Kokstad, (-DA), 4 Dec.

1928, Hutchinson 1798 (BOL, K); Ngele, (-DA), Mar. 1883, Tyson

1286 (SAM); 2 Jan. 1966, Strey 6365 (NU); 6 Jan. 1990, Abbott 4838

(NH); 13 Jan. 1990, Abbott 4925 (NH).

LESOTHO.— 2828 (Bethlehem): Leribe, (-CC), Dec. 1912, Diet-

erlen 6791 (SAM). 2927 (Maseru): Maseru, (-BD), 25 Jan. 1951,

Compton 22536 (NBG); between Molimo Nthuse and Blue Mtn Pass,

2 250 m, (-BD), 15 Feb. 1978, Schmitz 8095 (PRE); slopes behind

Malimo Nthuse Hotel, (-BD), 12 Jan. 1979, Hilliard & Burtt 12065

Bothalia 42,2 (2012)

159

(NU). 2929 (Underberg): Mokhotlong, (-AC), 25 Feb. 1949, Comp-

ton 21498 (NBG); 23 km from Mokhotlong on road to Sani Top, 2

250 m, (-AC), 12 Feb. 1987, KUlick 4578 (MO, NH); Sehlabaethebe

Reserve, 8 900' [2 700 m], (-CC), 3 Feb. 1975, Bayliss 1317 (PRE).

3028 (Matatiele): ridge between Orange River and Maqaba Peak, near

Quachasnek, (-BA), 13 Mar. 1936, Galpin s.n. BOL31667 (BOL).

NORTHERN CAPE. — 2723 (Kuruman): Carrington, (-CB), Apr.

1940, Esterhiiysen 2172 (BOL). 2724 (Taung): Klein Boetsap, (-CC),

1910, Pagan s.n. PRE54332 (PRE); W of Harz River near Taung,

(-DA), 12 Feb. 1948, Rodin 3646 (BOL, MO, PRE). 2816 (Oran-

jemund): Richtersveld, Ploegberg S of Khubus, (-DB), 3 Sept. 1977,

Oliver, Tolken & Venter 508 (MO, NBG, PRE). 2917 (Springbok):

O’kiep, (-DB), Sept.-Oct. 1926, Pillans 4983 (BOL). 2922 (Prieska):

Niekerkshoop, (-BB), 26 Nov. 1935, Bryant 1165 (BOL). 3017

(Hondeklipbaai): Bowesdorp, (-BB), Sept. 1941, Stokoe s.n. (MO,

PRE, SAM); Brakwater on Arakoop, (-BB), 25 Aug. 1977, Thompson

& le Roux 48 (MO. NBG); Grootvlei Pass, (-BB), 3 Sept. 1980, Le

Roux 2786 (BOL, NBG); Kamiesberg Pass, (-BB), 7 Sept. 1981, Van

Berkel 430 (NBG); 12 Sept. 1993, Strid & Strid 37769 (PRE); Skilpad

Wild Flower Reserve, (-BB), 10 Aug. 1993, Grobler 39 (PRE). 3018

(Kamiesberg): Kamiesberg, Leliefontein, (-AC), 16 Dec. \936, Adam-

son 1454 (PRE); Studer’s Pass, (-AC), 11 Sept. 2007, Snijman 2149

(NBG); Farm Damsland, (-AC), 29 Oct. 2007, Snijman 2213 (NBG);

Langkloof, N of Farm Doringkraal, (-AC), 23 Sept. 2010, Goldblalt

& Porter 135 73 (MO, NBG). 3119 (Calvinia): Nieuwoudtville Water-

fall, (-AC), Sept. 1930, La\’is s.n. BOL19622 (BOL); Nieuwoudtville

Reserve, (-AC), 7 Sept. 1983, Perry & Snijman 2312 (NBG, PRE);

13 Sept. 2010, Goldblatt & Porter 13526 (NBG); Oorlogskloof Nature

reserve, (-AC), 20 Sept. 1995, Pretorius 307 (NBG); between Oor-

logskloof and Papkuilsfontein, (-CA), Sept. 1939, Leipoldt 3076

(BOL); Calvinia, Farm Driefontein, (-DA), 23 Sept. 2009, Goldblatt,

Manning & Porter 13431 (MO, NBG). 3121 (Fraserburg): Platkop-

pies Farm, 45 km N of Williston, (-AA), 29 Mar. 1993, Germishuizen

6358 (PRE). 3122 (Loxton): 17 km from Loxton on road to Victoria

West, Taaibosfontein, (-BC), 13 May 1976, Thompson 3053 (NBG,

PRE). 3124 (Hanover): Richmond Dist., Elandspoort, (-CB), Oct.

1935, Thorne s.n. SAM51851 (SAM). 3221 (Merweville): Ezels Kom,

(-BA), 12 Apr. 1986, Shearing 1263 (PRE). 3222 (Beaufort West):

Karoo National Park, (-AB), 3 Jan. 1985, Shearing 866 (PRE). 3223

(Rietbron): Nelspoort, (-AA), without date, Pearson 2049 (SAM).

WESTERN CAPE.— 3118 (Vanrhynsdorp): Gifberg, (-DC), 15

Oct. 1953, Esterhiiysen 22132 (BOL); Gifberg Pass, (-DD), 11 Sept.

2009, Goldblatt & Porter 13319 (MO, NBG). 3217 (Vredenburg):

Witteklip, (-DD), 8 Sept. 2009, Goldblatt & Porter 13281 (MO, NBG,

PRE). 3218 (Clanwilliam): between Leipoldtville and Eland’s Bay,

(-AC), Oct. 1947, Zinn s.n. SAM63437 (SAM); Pakhuis Pass, (-BB),

4 Sept. 1948, Morris 20922 (BOL); 4 Sept. 1948, Compton 20922

(NBG); N7 S of Clanwilliam at Kransvlei turnoff, (-BB), 12 Sept.

2010, Goldblatt & Porter 13523 (MO, NBG); ± 4 km NE of Rede-

linghuys, (-BC), 9 Sept. 2009, Helme 6262 (NBG); Citrusdal, Mod-

derfontein, (-CB), 27 Aug. 1968, Hanekom 1163 (NBG, PRE). 3219

(Wuppertal): Brakfontein, (-AC), 20 Oct. 1983, Viviers 1245 (NBG);

Algeria Forest Station, (-AC), 20 Nov. 1996, Van Rooyen, Steyn & de

Villiers 253 (NBG); Matjiesrivier, (-AC), 6 Apr. 1944, Wagener 359

(NBG); Dasklip Pass, (-CC), 30 Sept. 1972, Thompson 1517 (NBG,

PRE). 3318 (Cape Town): Langebaan, Lynch point, (-AA), 5 Sept.

1971, Axelson 478 (NBG); near Darling, (-AD), Sept. 1905, Bolus

s.n. (BOL); hollow S of Strand Railway, Stickland, (-CD), Oct. 1932,

Acocks 959 (S); Camps Bay, (-CD), Oct. without year, Ecklon & Zey-

her s.n. (SAM); Table Mtn, (-CD), 29 Jul. 1846, Prior s.n. (PRE);

Sept. 1897, Thode 5828 (NBG); Lion’s Head, (-CD), 17 Oct. 1897,

Wolley Dod 3511 (BOL); Sept. 1913, Kensit & Teague s.n BOL45388

(BOL); Devil’s Peak, (-CD), Oct. 1877, Bolus 3844 (BOL); 31 Oct.

1915, Pillans 2808 (BOL); Table Mtn, Kasteelpoort, (-CD), without

date, Marloth 8999 (PRE); Malmesbury Dist., Helderfontein Farm,

(-DA), 11 Sept. 1979, Boucher 4684 (NBG, PRE); Langverwacht

above Kuil’s River, (-DC), 1 Oct. 1973, Oliver 4700 (NBG); Botlary-

berg, Farm Koopmanskloof, (-DD), 20 Sept. 1980, Beyers 86 (NBG);

above Jonkershoek, Haelkop, (-DD), 23 Oct. 1928, Gillett 1784 (BOL,

NBG); Jonkershoek, Bosboukloof, (-DD), May 1967, Kerfoot 5795

(NBG, PRE); Jonkershoek State Forest, (-DD), 31 Oct. 1975, Hay-

nes 1113 (NBG, PRE). 3319 (Worcester): Witzenberg, Inkruip, (-AA),

1 Oct. 1954, Esterhuysen 234219 (BOL); 5 km N of Tulbagh on way

to Winterhoek State Forest, (-AA), 8 Oct. 1981, Mauve & Hugo 59

(MO, NBG, PRE); Ceres Dist., Gydouw Pass, (-AB), 12 Aug. 1986,

Van Wyk 2530 (PRE); road from Hottentot’s Kloof to karoopoort, (-

BA), 29 Nov. 1908, Pearson 4831 (BOL); Dal Josafat Forest Reserve,

(-CA), 19 Oct. 1963, Taylor 5446 (NBG); Du Toil’s Peak, (-CA), 21

Dec. 1975, Esterhuysen 34165 (BOL); Fonteinjiesberg, (-CB), 1 Jan.

1976, Esterhuysen 34171 (BOL); Wemmershoek, (-CC), Jan. 1921,

Andreae 778 (PRE); Robertson Dist., Vrolijkheid Nature Reserve, (-

DD), 8 Apr. 1974, Theron & Students 3152 (PRE). 3320 (Montagu):

Montagu Div., Keurkloof, (-CC), 20 Sept. 1935, Lewis s.n. BOL31666

(BOL); Montagu-Barrydale Road, (-CD), Sept. 1923, Levyns 526

(BOL). 3321 (Ladimsith): Swartberg, Towerkop, (-AC), 16 Dec.

1956, Esterhuysen 26789 (BOL); Seweweekspoort, (-AD), 27 Dec.

1928, Andreae 1225 (PRE); Huis River Pass, (-BC), 31 July 1955,

Van Niekerk 536 (BOL); Meiringspoort, (-BC), Feb. 1932, Thorne

s.n. SAM50187 (SAM); Rooiberg, (-DA), 1 Nov. 1931, Compton 3827

(BOL). 3322 (Willowmore): Swartberg Pass, (-AC), Dec. 1904, Bolus

s.n. (BOL); Dec. 1943, Stokoe 9061 (BOL); Cango Valley, Bassonsrus,

(-AC), 4 Nov. 1974, Moffett 401 (PRE); Swartberg, Blouberg Peak,

(-AC), 29 Nov. 1987, Bean & Viviers 1987 (BOL); Wilderness, Fairy

Knowe, seashore dunes, (-DC), 13 Dec. 1928, Mogg 11896 (PRE);

Wilderness, (-DC), 14 Nov. 1952, Van Niekerk 196 (BOL); Ebb and

Flow Nature Reserve, Touw River Valley, (-DC), 19 Oct. 1971, Taylor

7996a (NBG, PRE). 3323 (Willowmore): Louterwater, (-CC), 18 Dec.

1933, Compton 45 73 (BOL); Herold’s Bay, (-CD), without date, Hugo

2118 (NBG); 17 Sept. 2010, Goldblatt & Porter 13546 (NBG). 3418

(Simonstown): Sandy Bay, (-AB), 26 Aug. 1978, Van Jaarsveld 3464

(MO, NBG); Constantiaberg, (-AB), 17 Dec. 1939, Compton 8209

(NBG); Hout Bay, (-AB), 16 Aug. 1928, Gillett 411 (NBG); Chap-

man’s Peak, (-AB), 7 Dec. 1943, Van Niekerk 469 (NBG, PRE); Nov.

1944, Le\yns 57806 (SAM); Silvermine, (-AB), 4 Oct. 1967, Bayliss

4006 (NBG); Kalk Bay Mtns, (-AB), 30 Oct. 1931, Adamson 2230

(BOL); 22 Nov. 1943, Lexyns 3553 (BOL); cliffs beyond Simonstown,

(-AB), 4 Oct. 1928, Hutchinson 646 (BOL, PRE); S of Simonstown,

above Froggy Pond, (-AB), 16 Nov. 1975, Esterhuysen 34107 (BOL);

Buffelsbaai, (-AD), 28 Nov. 1939, Adamson 2710 (BOL); Somerset

West, (-AB), 31 Oct. 1948, Parker 4378 (BOL, MO, NBG, S); Kogel

Bay, (-BD), 5 Sept, 1946, Parker 4111 (BOL, NBG); Rooi Els, (-BD),

15 Oct. 1971, Boucher 1670 (NBG); Pringle Bay, (-BD), 23 Dec.

1982, O'Callaghan 481 (NBG); Betty’s Bay, Jackass Penguin Col-

ony, (-BD), 16 Feb. 1997, Mucina 50359/1 (PRE). 3419 (Caledon):

Houhoek, (-AA), Apr. 1892, Guthrie 224 (NBG); Caledon, Warm-

baths Hotel, (-AB), 5 Oct. 1928, Gillett 1114 (NBG); 10 miles [16 km]

from Caledon on Bot River Road, (-AB), 17 Sept. 1938, Thorns s.n.

(NBG); Hermanns [cult, at Kirstenbosch], (-AC), Jan. 1932, Davis s.n.

NBG20/31 (BOL); Hermanns, cliffs opposite Marine Hotel, (-AC), 28

Aug. 2010, Manning 3304 (NBG); Buffeljagsbaai, (-DA/DC), 7 Sept.

1995, Paterson-Jones 617 (NBG). 3420 (Bredasdorp): Rietfontein,

20 km S Stormsvlei, (-AA), 23 Sept. 1982, Bayer 3071 (NBG); ± 25

miles [40 km] from Swellendam on Bredasdorp Road, (-AC), 30 Aug.

1962, Taylor 3847 (NBG); Windhoek, Potberg Nature Reserve, (-

AD), Jul./Aug. 1968, Van der Merwe 870 (PRE); 11 Oct. 1978, Burg-

ers 1243 (NBG). 3421 (Riversdale): near Kafir Kuils River, (-AB),

14 Oct. 1923, Muir 2916 (PRE); Stillbay rubbish dumps, (-AC), 9

Sept. 1978, Bohnen 4091 (NBG); Stillbay Rifle Range, (-AD), 2 Oct.

1978, Bohnen 4292 (NBG). 3422 (Mossel Bay): Goukamma Nature

Reserve, (-BB), 1969, Heinecken 245 (PRE); Buffalo Bay, (-BB), 26

Nov. 1955, Taylor 4908 (NBG). 3423 (Knysna): between Georgetown

[George] and Swellendam, (-AA), Jul. 1856, Castelnau 77 (BOL,

PRE); Bitou Bridge, (-AA), 15 Dec. 1941, Fourcade 5522B (BOL,

NBG); Knysna Heads, (-AA), 7 Nov. 1928, Gillett 2171 (BOL, NBG);

foot of Prince Alfred’s Pass, (-AA), Oct. 1932, Fourcade 4853 (BOL,

NBG); Keurboomstrand, (-AB), 14 Nov. 1949, Morris 433 (NBG); 25

Nov. 1955, Taylor 4917 (NBG); 17 Sept. 1956, Theron 2099 (PRE);

Ratels Bosch, (-BA), Aug. 1905, Fourcade 16 (BOL); Aug. 1908,

Fourcade 317 (BOL). 3424 (Humansdorp): Groot River, (-AA), Oct.

1942, Fourcade 5784 (BOL).

EASTERN CAPE. — 3027 (Lady Grey): Witteberg, Joubert’s Pass,

(-BC), 18 Jan. 1979, Hilliard & Burn 12206 (NU); Witteberg, Bed-

dgelert, (-DA), 2 Dec. 1981, Hilliard & Burn 14637 (E, NU); Ben

McDhui, (-DB), 9 Feb. 1983, Hilliard & Burn 16558 (E, K, NU).

3028 (Matatiele): Naude’s Nek, (-CA), 27 Dec. 1977, Bigalke 23

(NU); 13 Feb. 1983, Hilliard & Burn 16601 (E, NU). 3126 (Queens-

town): Jamestown, Vogelfontein Farm, (-BB), 16 Dec. 1942, Barker

2246 (NBG); Dordrecht, (-DB), Apr. 1929, Smuts s.n. (NBG); 22

Mar. 1964, Bayliss 2119 (NBG). 3127 (Lady Frere): Maclear, Baster-

voetpad. Farm Snowy Side, (-BB), 17 Nov. 1993, Bester 1713 (NH).

3128 (Umtata): Farm Comarty ± 18 km W of Ugie, (-AA), 19 Apr.

1994, Bester 2712 (NH); Libode, Misty Mount, S of Port St. Johns,

(-DB), 18 Nov. 1991, Cloete 1344 (NH). 3224 (Graaf-Reinet): ‘Water-

furrows’ above Graaf-Reinet, (-BA), Nov. 1865, without collector

BOL74044 (BOL). 3226 (Fort Beaufort): Pluto’s Vale, (-BA), 29 Oct.

1964, Bayliss 2481 (NBG); Elandsberg, (-BD), 17 Jan. 1986, Cadman,

Edwards & Norris 3224 (NU); Hogsback, (-DB), May 1932, Leighton

160

Bothalia 42,2 (2012)

s.n. BOL31669 (BOL); June 1961, Bokelmami 6 (NBG). 3227 (Stut-

terheim): Kleinemonde, (-CA), ?Nov. 1953, Tavlor 4306 (NBG);

King William’s Town, (-CD), Jan. 1894, Sim 1613 (NU). 3228 (But-

terworth): Kei Mouth, (-CB), Aug. 1891, Flanagan 880 (SAM). 3325

(Port Elizabeth): Walmer, (-DC), Sept. 1916, Paterson 3321 (NBG,

PRE); Redhouse, (-DC), Sept. 1914, Paterson 272 (BOL); Baakens

River Valley, Fern Glen, (-DC), 14 Nov. 1974, Olivier 1241 (MO,

NBG). 3326 (Grahamstown): Martindale, Kap River, (-BD), 23 Dec.

1956, Taylor 5180 (NBG). 3425 (Skoenmakerskop): Sea View, (-AB),

18 Sept. 2010, Goldblatt 13549 (NBG); Sardinia Bay, (-BA), 18 Sept.

2010, Goldblatt 13552 (NBG); coast near Cape Recife, (-BA), 18

Sept. 2010, Goldblatt & Porter 13553 (NBG),

2b. subsp. polyantha J.C. Manning & Goldblatt,

subsp. nov.

TYPE. — KwaZulu-Natal 2930 (Pietermaritzburg):

Krantzkloof, (-DD), Oct. 1921, Haygarth 138 (NBG,

holo.; NH, iso.).

Stems erect. Cymes relatively many-flowered, with

tertiary and often quartemary branching, fonning a flat-

topped, corymbiform synflorescence. Calyx 12-15(-20)

mm long. Capsules broadly ovoid, 10-15 mm long, 4-6

X longer than carpophore. Carpophore 2-4 mm long.

Distribution and ecology’: restricted to the coastal

and near-inland parts of KwaZulu-Natal and Swaziland

between 550-1 000 m, with a single early record from

Barberton (Figure 8), occurring along thicket and forest

margins and in marshy places.

Diagnosis: distinguished from the typical form by

the erect stems and closely branched, flat-topped inflo-

rescence, with tertiary and often quartemary branching,

short calyx, mostly 12-15(-20) mm long, and short,

broadly ovoid capsules, 10-15 mm long with very

short carpophore, 2-A mm long. This form grades into

the typical subspecies in the KwaZulu-Natal Midlands,

between Pietermaritzburg and Vryheid.

Additional specimens

MPUMALANGA. — 2531 (Komatipoort): Barberton, (-CC), Dec.

1916, Port 5467 (PRE).

SWAZILAND.- 2631 (Mbabane): Mbabane, Forbes Reef, (-AC),

29 Nov. 1963, Compton 31797 (NBG); Black Mbuluzi Valley, 3500'

[I 060 m], 9 Dec. 1958, Compton 28443 (NBG); 9 Dec. 1964, Comp-

ton 32183 (NBG); Gobolo, (-AC), 23 Nov. 1962, DIamini s.n. (PRE);

Fllatikulu, (-CD), Dec. 1910, Stewart s.n. SAM2523 (SAM); 23 Nov.

1959, Compton 29502 (NBG).

KWAZULU-NATAL. — 2731 (Louwsburg): Itala Nature Reserve,

(-CB), 1 1 Dec. 1975, Hilliard & Biirtt 8551 (E, K, NU, PRE, S); Non-

goma, (-DC), 2 Dec. 1943, Gerstner 4666 (NBG). 2831 (Nkandia):

Eshowe, Reservoir March, (-CD), 9 Nov. 1949, Lawn 1296 (NFI).

2832 (Mtubaluba): Dukuduku, (-AC), 25 Nov. 1965, Strey 6099 (NU,

PRE). 2930 (Pietermaritzburg): ± 25 km NNE Howick, Twin Falls,

(-AD), 19 Nov. 1987, Grove 122 (NU); Baynesfield, Oldfield Farm,

(-CD), Oct. 1989, Edwards 602 (NU); Hammarsdale, 590 m, (-DA),

27 Jun. 1995, Ward 13189 (NH, NU, PRE); Botha’s Hill, (-DB), 10

Dec. 1913, Medley Wood 12387 (NH, NU). 3029 (Kokstad): Harding,

Rooi Vaal, (-DB), 4 Jan. 1957, Taylor 5275 (NBG). 3030 (Port Shep-

stone): Alexandra Dist., Umgay [Umgai], (-AD), 9 Oct. 1909, Rndatis

522 (NBG); Rudatis 741 (PRE); Campbellton-Dumisa, (-AD), 13 Nov.

1913, Rndatis 2030 (NBG); near Southport, (-DA), 13 Nov. 1974,

Nicholson 1506 (PRE).

3. Silene saldanhensis J.C. Manning & Goldblatt, sp.

nov.

TYPE. — Western Cape 3317 (Saldanha); Saldanha

harbour, SW-facing slopes on ridge at beginning of

breakwater, (-BB), calcareous sand, 19 Sept. 2011,

Goldblatt & Manning 13646 (NBG, holo.; K, MO, PRE,

S, iso.).

Compact or cushion-forming perennial, 30-60 cm,

branching at base with multiple crowns; stems erect or

decumbent, 2-4 mm diam., with shoots in lower axils,

densely pubemlous with gland-tipped hairs mixed with

scattered, short, acute eglandular hairs 0.25-0.50 mm

long, inflorescence axis often especially densely puber-

ulous. Leaves fleshy, densely pubemlous with mixture

of erect, gland-tipped hairs and acute, eglandular hairs,

ciliate at extreme base with longer hairs to 1 mm long,

with evident side veins, basal leaves oblaneeolate, up to

100 X 35 mm; stem leaves spreading, lanceolate, 35-50

X 12-18 mm, recurved apically, margins weakly revo-

lute and sometimes undulate. Inflorescence a lax, few-

flowered, assymetrical, paniculate cyme, axis 1.5-2. 5

mm diam. below primary flower; bracts similar to upper

leaves but smaller, spreading-recurved; primary pedicels

10-20 mm long, elongating to 35 mm in fruit, second-

ary pedicels 10-20 mm long. Flowers half-nodding,

nocturnal and faintly gardenia-scented at night. Calyx

um-shaped, 17-20 mm long, with ± prominent, thick-

ened ribs, 10-ribbed, with 1-3 reticulate veins in distal

half, densely pubemlous with erect, gland-tipped hairs,

lobes triangular, 3^ mm long, densely glandular-ciliol-

ate. Petals mauve, sometimes with darker corona, limb

spreading, broadly cuneate, ± overlapping with adjacent

petals, 13-18 x 10-19 mm, bifid ± one third to halfway,

sometimes crenulate distally, one or both margins some-

times with small sub-basal tooth, claw strap-shaped,

15-20 X 3*M mm, auriculate apically, glabrous or

sparsely pubescent along abaxial midline and margins,

coronal scales suberect, 1. 5-2.0 mm. Stamen filaments

subequal, 15-18 mm long, reaching to top of claws or

shortly exserted but not exceeding corona; anthers ± 2

mm long, not exserted beyond corona. Ovary nan'owly

ellipsoid, 6 mm long, stipe ± 1mm long; styles ulti-

mately to 8 mm long, reaching top of corona. Capsule

ovoid, 17 X 6. 5-8.0 mm, ± 5 times longer than carpo-

phore, minutely granular; carpophore 3 mm long, retro-

sely scabridulous. Seeds 1.0-1. 2 m diam., subcircular,

charcoal grey, with scattered whitish or translucent testal

cells, face flat or concave, back flat, testal cells concen-

trically colliculate. Flowering time: Aug.-Oct. Figures

2C, D, 9.

Distribution and ecology: largely restricted to the

entrance to Saldanha Bay, where it is known from the

breakwater and from across the lagoon in Postberg

Reserve (Figure 10), but also known from shortly inland

on the Farm Waterboerskraal east of Hopefield. The spe-

cies is restricted to calcareous sands and consolidated

calcrete dunes.

Diagnosis: a compact or cushion-forming perennial

with erect or decumbent stems, distinguished by the

mauve flowers with broad, overlapping petal limbs. The

stamens and styles are included or shortly exserted but

never protrude beyond the corona. Silene saldanhensis

is probably most closely allied to the carmine-flowered

S. ornata (Figure 1 1 ), which it resembles in stem ves-

titure and in the presence of scattered translucent testal

cells in the seeds, but is distinguished by its broader,

mauve flowers, colliculate seeds (Figures 2C, D), and in

its ecology. Both species have stems with a mix of glan-

Bothalia 42,2 (2012)

161

FIGURE 9. — Silene saldanhensis, Saldanha, Goldblatt & Manning 13646. A, flowering stem; B, stem segment; C, calyx laid out; D, petal; E,

androecium laid out; F, gynoecium; G, capsules (left hand illustration before dehiscence with caljoc removed; central after dehiscence, right

hand after dehiscence with calyx removed); H, seed. Scale bar: A, G, 10 mm; B, 1 mm; C, 5 mm; D-F, 2.5 mm; H, 0.5. Artist: John Manning.

dular hairs and short, scattered eglandular hairs (Figures

9B, IIB) but S. ornata is restricted to rocky habitats,

and typically has a straggling growth form, even in very

exposed situations, with the stems sprawling among

bushes or over the rock surface. It typically has slightly

smaller flowers, with petal claws ±15 mm long and the

limbs not overlapping, and tubereulate seeds (Figures

2E, F). Silene saldanhensis favours deep, loose sands or

compaeted dunes, forming neat cushions with the stems

either decumbent or erect. It has mostly larger flowers

with the claws 15-20 mm long and broad, often overlap-

ping petals, and colliculate seeds, which are distinctive

162

Bothalia 42,2 (2012)

FIGURE 10. — Distribution of Silene saldanhensis, o; S. ornata, •; S.

rigens. A; S. mundiana, ▲ .

among the southern African members of sect. Elisan-

the in lacking papillae or tubercules on the testal cells.

The mix of glandular and short, eglandular hairs on the

stems, and broad, overlapping petals, with included sta-

mens and styles separate S. saldanhensis from coastal

forms of S. undulata with a short calyx and carpophore.

Conservation note: the restricted occurrence of the

species, the low number of plants and development

at Saldanha render the species threatened. The Post-

berg population is protected as part of the West Coast

National Park.

Additional specimens

WESTERN CAPE.— 3317 (Saldanha): West Coast National Park,

Kraalbaai, (-BB), 13 Sept. 2011, Manning & Goldblatt 3339 (NBG).

3318 (Cape Town): Hopefield, Waterboerskraal Farm, (-AB), 3 Get.

\974, Hugo s.n. (NBG).

4. Silene ornata Aiton, Hortus kewensis 2: 96 (1789);

Sond.: 126 (1860). Melandrium ornatum (Ait.) Asch. ex

Rohrb.: 233 (1869). Type: South Africa, without precise

locality, cultivated at Kew in 1774 from seed introduced

by Francis Masson, Masson s.n. (BM000573045, holo.!;

BM000573046, iso.!). Illustrated in Curtis’s Botanical

Magazine 1 1 : t. 382 ( 1 797).

Silene hellidifolia sensu Thunb.: 81 (1794), non. Juss.

ex Jacq. (1777).

Perennial to 1 m, well branched with branches

ascending at ± 45 degrees, or plants compact and almost

cushion-forming in exposed situations; stems sprawl-

ing among vegetation or on rock surface, 1.5-2. 5 mm

diam., densely puberulous with spreading, gland-tipped

hairs mixed with equally short, acute hairs 0.25-0.50

mm long, inflorescence axis often especially densely

puberulous. Leaves mostly ± cauline, spreading, lower

leaves oblanceolate-spathulate, sub-petiolate; stem

leaves lanceolate to oblanceolate, mostly 50-80 x 10-20

mm, acute, base tapering or cuneate, margins plane or

undulate, puberulous with mixture of erect, gland-tipped

hairs and acute, eglandular hairs, ciliate at extreme base

with longer hairs to 1 mm long, with evident side veins.

Inflorescence a lax, few-flowered, assymetrical, pan-

iculate cyme, axis 1.0-1. 5 mm diam. below primary

flower; bracts similar to upper leaves but smaller, sub-

equal, spreading-recurved; primary pedicels 10^0 mm

long, secondary pedicels 6-18 mm long. Flowers ±

horizontally spreading on pedicels flexed sharply api-

cally, ± open day and night, unscented. Calyx narrowly

um-shaped in flower, 19-21 mm long, 10-ribbed, with

1-3 reticulate veins in distal half, densely puberulous

with erect, gland-tipped hairs, lobes narrowly triangu-

lar, 3-5 mm long, densely glandular-ciliolate. Petals

deep carmine, limb spreading, cuneate, not overlap-

ping with adjacent petals, (7-) 10-1 5 x (7-) 12-1 5 mm,

bifid halfway to two thirds, claw strap-shaped, ± 15

X 3 mm, auriculate apically, coronal scales suberect,

2. 0-2. 5 mm long. Stamen filaments subequal, 14—15

mm long, reaching to just below top of claws; anthers ±

2 mm long, reaching top of claws, not exserted beyond

corona. Ovary narrowly pyriform, shortly stipitate, ±

7 mm long, stipe ± 1mm long; styles 7-12 mm long,

included or exserted up to 4 mm beyond petal claw.

Capsule ovoid, 15-18 x 6-7 mm, 4—5 x longer than car-

pophore, minutely granular; carpophore ± 4 mm long,

glabrescent. Seeds 1.0-1. 2 mm diam., subcircular with

hilum recessed, grey or reddish brown, with scattered

translucent cells, face flat or concave, back flat, testal

cells radial-concentric, tuberculate. Flowering time: (late

Aug.) Sept.-Oct. Figures 2E, F, 11).

Distribution and ecology, a local endemic of the

limestone hills flanking the mouth of Saldanha Bay

(Figure 10), occurring on rocky slopes among coastal

scrub. Plants mostly occur on the slopes or summit of

hills, with the annual stems straggling through surround-

ing shrubs, but a population at Stony Head thrives on

exposed limestone pavement directly facing the ocean.

The plants here are very compact and almost cushion-

forming, with the flowering stems sprawling along the

rock sheets.

Diagnosis and relationships'. Silene ornata is readily

recognized by the unscented, deep carmine flowers that

remain open thoughout the day and night. The branch-

ing of the flowering stems is largely asymmetrical, and

stems and branches are densely puberulous, with a mix

of short, glandular hairs and non-glandular hairs. The

stamens are always included and the styles mostly so but

in some plants they are exserted up to 4 mm beyond the

petal claws. This is the form that is illustrated in Cur-

tis’s Botanical Magazine 11: t. 382 (1797). Other species

in sect. Elisanthe have nocturnal, white to pale pink or

mauve flowers that are fragrant at night. Silene ornata is

evidently most closely allied to S. saldanhensis, another

local endemic of Saldanha Bay, which shares a similar

stem vestiture but which is restricted to calcareous sands

and consolidated dunes, has slightly larger, mauve flow-

ers with broader tepals, a more tufted habit, and collicu-

late seeds.

The species, named by Aiton (1789) for its strik-

ing, dark red flowers, was introduced to Kew by Fran-

cis Masson in 1775. Although the exact provenance of

Masson’s material is not given, Silene ornata is a nar-

row endemic of Saldanha Bay, which Masson vis-

ited for five days in September 1773 in the company

of Carl Thunberg (Glen & Germishuizen 2010), and it

Bothalia 42,2 (2012)

163

FIGURE 1 1 . — Silene ornata, Saldan-

ha, Goldblatt & Porter 13254.

A, flowering stems; B, stem

segment; C, calyx laid out; D,

petal; E, capsules (left hand

illustration before dehiscence

with calyx removed; central

after dehiscence, right hand

after dehiscence with calyx

removed); F, seed. Scale bar:

A, E, 10 mm; B, 1 mm; C, 5

mm; D, 2.5 mm; F, 0.5. Artist:

John Manning.

is probable that they collected the species at this time.

Thunberg (1794) mistakenly associated his collection

{UPS-THUNB 10705) with the bright pink-flowered

Mediterranean S. bellidifolia, as correctly realized by

Sonder ( 1 860). Masson’s brief on the trip was to collect

seeds, cuttings and roots for cultivation at Kew and he

certainly succeeded with S. ornata. Readily propagated

by cuttings, the species was widely cultivated in Eng-

land by 1797, when it was illustrated in Curtis’s Botani-

cal Magazine (Curtis 1797) but has since disappeared

from cultivation.

Conservation note', the restricted occurrence of the

species, the low number of plants and the encroaching

urbanisation at Saldanha render the species threatened.

The Postberg population is protected as part of the West

Coast National Park.

Additional specimens

WESTERN CAPE. — 3217 (Vredenburg): limestone hill north

of Saldanha, (-DD), 17 Sept. 1976, Goldblatt 4114 (MO); hill N of

Saldanha, (-DD), 250' [76 m], 24 Sept. 2009, Goldblatt & Porter

13254 (K, MO, NBG, PRE). 3317 (Saldanha): Hoedjies Bay, (-BB),

Sept. 1905, Bolus 12614 (BOL); Postberg, (-BB), 8 Sept. 1957, Lewis

52 73 (NBG); 30 Aug. 1980, Bond 1715 (MO, NBG); hills between

Postberg and Donkergat, (-BB), 9 Sept. 1966, Rourke 576 (NBG);

Konstabelkop and Postberg hills, (-BB), 13 Sept. 2011, Manning &

Goldblatt 3340 (NBG); SAS Saldanha, Bomgat, (-BB), 15 Sept. 2011,

CR15241 (NBG). Without precise locality: hort. Kirstenbosch, Aug.

1937, without collector BOL31664 (BOL).

164

Bothalia 42,2 (2012)

5. Silene rigtm J.C. Manning & Goldblatt, sp. nov.

TYPE. — Western Cape 3218 (Clanwilliam): Lange-

baanweg. Fossil Park road, (-CC), 127' [38.4 m], 8 Sept.

2009, Goldblatt & Porter 13291 (NBG, holo.; MO,

PRE, iso.).

Silene noctiflora sensu Thunb., in part [excluding ref-

erences to Table Mtn and Swartland]: 81 (1794), non. L.

(1753).

Tufted perennial, 0.5-1. 5 m, producing one or two

new vegetative shoots from basal axils at end of flow-

ering season; rootstock a taproot; stems stiffly erect,

unbranched except in inflorescence, 3-5 mm diam.,

densely puberulous with spreading, gland-tipped hairs

0.2-0. 5 mm long. Leaves mostly basal, suberect or

spreading, lower leaves oblanceolate-spathulate, sub-

petiolate, 100-140 x 15-25 mm, dead or dying at flow-

ering, stem leaves elliptical to oblanceolate, mostly

60-100 X 10-20 mm, acute, base tapering, margins

plane or undulate, pubescent with mixture of erect,

gland-tipped hairs and acute, eglandular hairs when

young but apparently entirely eglandular at maturity,

ciliate at extreme base with longer hairs to 1 mm long,

with evident side veins. Inflorescence a compact, sym-

metrically branched cyme with flowers in triads, axis

1. 5-3.0 mm diam. below primary flower; bracts similar

to upper leaves but smaller, subequal, suberect; primary

pedicels 10^0 mm long, secondary pedicels 10-20 mm

long. Flowers suberect or half-nodding, nocturnal, jas-

mine-scented or unscented. Calyx narrowly um-shaped

in flower, 18-22 mm long, 10-ribbed, with 1^ reticu-

late veins in distal half, densely puberulous with erect,

gland-tipped hairs, lobes narrowly triangular, 3-6 mm

long, densely glandular-ciliolate, splitting almost to base

to expose mature capsule. Petals whitish or pale dingy

pink to mauve, limb spreading, cuneate, not overlap-

ping with adjacent petals, 8-10 x 6-8 mm, bifid ± half-

way, claw strap-shaped, 16-18 x 2.5 mm, auriculate

apically, coronal scales suberect, ± 1 mm long. Stamen

filaments unequal, shorter series 9-10 mm long, longer

series 12-14 mm long, reaching to 4 mm below top of

claws; anthers ± 2 mm long, not reaching top of claws,

included. Ovary narrowly pyriform, shortly stipitate,

5-6 mm long, stipe ± 0.25 mm long; styles ± 5 mm long,

included, ultimately barely exserted. Capsule broadly

ovoid, 15-20 X 9-10 mm, ± 6 x longer than carpophore,

minutely granular; carpophore 2. 5-3.0 mm long, gla-

brescent. Seeds 1.2-1. 5 mm diam., reniform with hilum

recessed, face concave, back flat, reddish brown, testal

cells radial-concentric, tuberculate on face but echinate

along shoulders and back. Flowering time: Sept.-Oct.

Figures 2G, H, 12.

Distribution and ecology, restricted to the coastal

forelands of the southwestern Cape, from Saldanha Bay

along the West Coast to the Cape Flats, and at Vermont

near Hermanns (Figure 10), occurring in deep, calcare-

ous sands in strandveld thicket.

Diagnosis and relationships: although overlooked as

a distinct species until now, Silene rigens is readily dis-

tinguished in sect. Elisanthe by its characteristic growth

form and stem vestiture, flowers, fruits, and seeds. The

stems are tall, stiffly erect, and largely unbranched

except in the inflorescence, where the branching is sym-

metrical and the branches relatively short, resulting in

distinctive, condensed triads of flowers and fruits. Flow-

ering stems are strictly annual, dying off at the end of

the season, to be replaced by axillary shoots from the

base. The stem vestiture comprises only gland-tipped

hairs, without an admixture of eglandular, acute hairs.

Individual flowers are relatively small for the section,

with white, dull pink or light mauve petals, the claws

8-10 mm long, and the anthers and styles ultimately

barely exserted. The capsules, which are unusually

broadly um-shaped at maturity, 15-18 x 9-10 mm, and

with a very short carpophore ±2.5 mm long, cannot be

fully accommodated within the calyx, which splits lon-

gitudinally at least midway but often to the base. The

distinctly echinate seeds (Figures 2G, H) are unique

among the southern African species of sect. Elisanthe,

which are otherwise colliculate or tuberculate.

Although known for almost 260 years, since the first

specimens were collected by Carl Thunberg during his

visit to the West Coast in 1773-1774, Silene rigens has

only now been recognized as a distinct species. Thun-

berg (1794) mistakenly associated his material (UPS-

THUNB10774 & 10775) with the European species S.

noctiflora L. and Sender (1860) subsequently treated

them under S. undulata. More recent collections of the

species have been variously misidentified as S. bellidi-

oides, S. capensis, and S. ornata.

Conservation note: the restricted occurrence of the

species and its local extinction on the Cape Flats render

it vulnerable, particualrly as most populations occur in

areas undergoing intensive development for housing,

industry and recreational activity.

Additional specimens

WESTERN CAPE.— 3317 (Saldanha): Hoedjies Bay, (-BB), Sept.

1905, Bolus 1 261 3 (BOL). 3318 (Cape Town): Langebaan Peninsula,

Stofbergsfontein, (-AA), 28 Nov. 1975 [sterile], Bucher 2963 (NBG);

Elandsfontein West, (-AA), 5 Oct. 1977, 500' [150 m], Thompson!

3525 (NBG); R27 to Langebaanweg, (-AB), 20 Sept. 2009, Goldblatt,

Manning & Porter 13409B (MO, NBG); Vygevallei farm, (-AD), 24

Sept. 1996, Low 2801 (NBG); Duynefontein (Koeberg), (-CB), 31

Oct. 1986, Bosenberg & Rutherford 177 (NBG); Duynefontein, (-CB),

30 Sept. 2010 [fruiting], Goldblatt & Manning 13599 (MO, NBG);

Melkbosstrand, (-CB), 8 Sept. 1940, Compton 9329 (NBG); Paarden

Eliland, (-CD), 19 Sept. 1942, Compton 13720 (NBG); damp hollow

near Strand Line, Stickland, (-CD), 23 Oct. 1932, Acocks 1049 (S);

Belville, Cape Flats Nature Reserve, University of Western Cape, (-

DC), 5 Aug. 1978, Low 578 (NBG); 31 Aug. 1978, Low 624 (NBG).

3418 (Simonstown): Cape Flats, Swartklip, along road to military hut

in E portion, (-BA), 22 Sept. 1972, Taylor 8211 (NBG, PRE). 3419

(Caledon): Vermont, near the sea, deep sand, (-AC), 4 Sept. 1986, S.

Williams 1163 (MO).

Silene sect. Fruticulosae (Willk.) Chowdhuri [as

(Rohrb.) Chowdhuri], Notes from the Royal Botanic

Garden, Edinburgh 22: 246 (1957). Silene [unranked]

Fruticulosae Willk., Icon. Descr. PI. Nov. 1: 73 (1854).

Silene ser. Fruticulosae (Willk.) Rohrb., App. Alt. Ind.

Sem. Hort. Bot. Berol. 1867: 2 (1867). Lectotype: Silene

ciliata Pourr., designated by Greuter: 573 (1995b).

[Chowdhuri’s (1957) lectotypification of the section

against S. burchellii Otth is treated as an error by Greu-

ter (1995b)].

Perennials or geophytes with woody or tuberous root-

stoek, eglandular pubescent. Leaves linear to oblaneeo-

Bothalia 42,2 (2012)

165

FIGURE 12. — Silene rigens, Lange-

baanweg, Goldblatt & Por-

ter 13291. A, base of plant;

B, flowering stem; C, stem

segment; D, calyx laid out;

E, petal; F, androecium laid

out; G, gynoecium; H, cap-

sules (left hand illustration

before dehiscence with calyx

removed; central before

dehiscence, right hand after

dehiscence); I, seed. Scale

bar: A, B, H, 10 mm; C, 1

mm; D, 5 mm; E-G, 2.5 mm;

1, 0.5. Artist: John Manning.

late, lacking apparent lateral veins; bracts herbaceous.

Flowers in monochasia with axis simple or forked

below, spreading or ± erect, vespertine; anthophore

well-developed, pubescent. Calyx pubescent, clavate,

10-veined, without conspicuous anatomosing venation,

lobes ovate, obtuse. Petals: claw glabrous or pubescent

along veins, weakly auriculate, limb bifid with linear

lobes; coronal scales present. Ovary partially septate.

Seeds reniform with deep hilar notch, flanks flat or

somewhat concave and smooth or striate, back deeply

and narrowly grooved between two undulate peripheral

wings.

± 12 spp., S Europe, Africa.

6. Silene burchellii Otth in DC., Prodromus syste-

matis naturalis regni vegetabilis 1: 374 (1824); Sond.;

128 (1860). Type: South Africa, [Western Cape],

kloof between Lion’s Head and Table Mt., on the ride

towards Camps Bay, without date [probably 1810/1811],

Burchell 271 (G-DC, holo.; K,!, PR£!, iso.). [Although

authorship of the species has been attributed either to

‘Otth ex DC.’ or to ‘DC.’ alone, De Candolle’s note

(Candolle 1824: 367) at the foot of the generic treatment

makes it clear that the author of the account is Adolphe

Otth, and new names published there should thus be

attributed to Otth alone.]

Geophytic perennial or suffrutex, mostly 10-50 cm

tall; tuber parsnip-like or ovoid; stems decumbent or

erect, ± suffrutescent below, basal woody portion 1-2

mm diam. but 0.5-1. 5 mm diam. at base of inflores-

cence, ± densely puberulous-scabridulous with short,

deflexed or spreading, acute hairs, rarely sub-glabrous.

166

Bothalia 42,2 (2012)

Leaves cauline but concentrated in lower part of stem,

± abruptly separated from inflorescence, lower some-

times dry or withered, linear to obovate, (10-) 15-60 x

l-15(-22) mm, obtuse, apiculate or weakly uncinate,

base narrowed, ± adpressed-puberulous, usually more

densely so abaxially, rarely glabrous on adaxial or both

surfaces, hairs acute, margins ciliate towards base with

longer, straggling hairs to 1 mm long, without evident

side veins. Inflorescence a lax or dense, (2)3-1 2-flow-

ered monochasium terminating main stems, well sepa-

rated from foliage, simple or with 1(2) branches from

base, rachis 0. 5-1.0 mm diam.; bracts smaller than

leaves, unequal to subequal, oblong or lanceolate to sub-

ulate, puberulous with appressed, acute hairs, margins

densely ciliolate; pedicels 3-10(-15) mm long, rarely

up to 20 mm long in fruit. Calyx clavate in flower, (10-

)12-18(-25) X 2.5^ mm at anthesis, slightly upcurved

distal to carpophore when > ± 15 mm long, equally

10-veined, adpressed-puberulous, lobes ovate or trian-

gular, ± 2-A mm long, densely ciliolate. Flowers ± pat-

ent or suberect at anthesis, nocturnal, coconut-scented

or more medicinal and acrid. Petals white to pale yel-

lowish, pale pink or mauve, with darker green or maroon

reverse, claw 7-8(-10) mm long, rarely exserted up to

6 mm beyond calyx, narrowed below, without evident

auricles, pubescent along midrib and veins on outer sur-

face, limb bifid, 3-7 x 2-5 mm, coronal scales 0.2-0. 8

mm long. Stamen filaments slightly unequal, 6-10 mm

long, shorter series shortly included, longer series reach-

ing top of claw or exserted up to 2 mm. Ovary ellipsoid,

± 4 mm long; styles 4-6 mm long, exserted up to 3 mm.

Capsule ovoid, 8-10 x 5-7 rnm, slightly longer than to

slightly shorter than carpophore, smooth; carpophore

4-12 mm long, pubescent. Seeds 1.0-1. 5 mm, reniform

with deep hilar notch, flanks flat or somewhat concave,

back deeply and narrowly grooved between two undu-

late wings, reddish brown, testa striate. Flowering time:

Aug.-Nov. in the winter rainfall region; mainly Oct-

May in the summer rainfall region but almost year-round

along the coast. Figures 3A-C, 13.

Distribution and ecology: widely distributed through

the more temperate parts of southern Africa but absent

from the arid interior, through eastern Africa northwards

to the highlands of Sudan and Ethiopia, and extending

across the Red Sea to Arabia (Figure 13). Silene burchel-

lii favours loamy or fine-grained clay soils, often in

rocky or stony places, from near sea level up to over

4 000 m. In southern Africa it is mostly a grassland forb

but in the winter rainfall region occurs in open, mostly

drier, fynbos and succulent shrubland.

Diagnosis and relationships: Silene burchellii is dis-

tinguished from the other southern African members of

sect. Fruticulosae by its mostly erect or suberect stems,

0.5-1. 5 mm diam. at the base of the inflorescence, and

mostly linear to oblanceolate, rarely obovate leaves,

(10-)15-60 X 1-16 mm. The leaves are ± concentrated

in the lower part of the stem and well separated from the

inflorescence by a distinct, leafless upper part ±100-150

mm long bearing one or two pairs of bract-like leaves.

The calyx is very variable in length, 10-25 mm long,

and is often slightly upcurved. The carpophore ranges

from 4-12 mm long, varying from slightly longer than

to shorter than the um-shaped capsule, 8-10 x 5-7 mm.

Silene burchellii is sometimes confused with S. aethi-

opica in the Western Cape, but is distinguished from

this annual species by its perennial, geophytic habit, and

also never develops the highly branched growth form

evident in well-grown plants of S. aethiopica. Although

the tuber is seldom present on herbarium specimens, the

relatively thick stem, terminating abruptly where it has

been broken from the tuber, is characteristic. The two

species are also ecologically separated, with S. aethio-

pica mainly coastal in deep sandy soils and S. burchellii

essentially an inland species on finer-grained clay soils.

Along the eastern seaboard, especially the Wild Coast

and KwaZulu-Natal, broad-leaved S. burchellii subsp.

multiflora can be confused with S. crassifolia subsp.

primuliflora. The latter can usually be identified by its

prostrate or decumbent, often branching stems, mostly

1 .5-2.0 mm diam. at the base of the inflorescence, leath-

ery or thick-textured leaves, somewhat expanded and

strongly pleated calyx, mostly larger coronal scales,

0. 8-1.0 mm long, and broadly um-shaped to subglobose

capsules, (8-) 10- 12 x 7_io mm. Its strictly coastal dis-

tribution, on sandy foredunes, is also diagnostic.

As currently circumscribed, Silene burchellii is the

most widely distributed of the African species, ranging

through the length of the continent, from Arabia and the

uplands of Sudan and Ethiopia southwards though East

Africa to the Cape Peninsula (Wickens 1976). Although

highly variable in leaf shape and width, development of

indumentum, number of flowers, and in calyx length, the

species has resisted all attempts at segregation into more

finely circumscribed taxa. The variation is often broadly

correlated with geography and/or ecology but mor-

phological patterns are confounded by the existence of

numerous intermediates. The situation was summarised

several decades ago by Turrill (1954), who concluded

that it was reasonable to assume that the variation in S.

burchellii is associated both with genetic and with envi-

ronmental differences. This opinion has not altered since

then, and Hedberg (1954, 1957) was unable to maintain

any afro-alpine inffaspecific taxa on the morphologi-

cal evidence without supporting evidence from further

breeding experiments.

Despite the intractability of the species to satisfac-

tory subdivision, workers on the tropical Africa mate-

rial have consistently commented on the evident differ-

ences between the narrow-leaved tropical plants and the

type of the species from the southwestern Cape, which

is characterized by ± prostrate or decumbent stems and

broad, oblanceolate to obovate leaves (Turrill 1954,

1956a; Wild 1961; Gilbert 2000). There is a general feel-

ing that the tropical African representatives should be

distinguished at some level from all or at least some of

the southern African material, and Wild (1961) accord-

ingly treated all collections from the Flora zambesi-

aca area as var. angustifolia Sond. (1860) to separate it

from typical var. burchellii from South Africa. With this

signal exception, Sonder’s (1860) early attempt at seg-

regating the South African material of the species into

four varieties, based largely on leaf shape and growth

form, has not been implemented. The recent treatment

of the species in Ethiopia and Eritrea by Gilbert (2000)

avoided all recognition of formal varieties in favour of

five informal ‘forms’.

Bothalia 42,2 (2012)

167

FIGURE 13. — Silene burchellii: A,

subsp. burchellii. Signal Hill,

no voucher; B, subsp. pilosel-

lifolia, Kamiesberg, Goldblatt

& Porter 13574. Scale bar: 10

mm. Artist: John Manning.

168

Bothalia42,2 (2012)

The southern African collections display exten-

sive variation in vegetative and floral morphology but,

as with the tropical African material, this is ± continu-

ous, with no clear separation between the extremes, and

we are unable to recognize more than a single species.

There are, however, some general correllations between

morphology and geography that we interpret as demon-

strating some level of genetic differentiation within the

species. Subsuming all of this variation under a single

name obscures its existence and we therefore propose

the recognition of four subspecies to highlight the most

distinctive variants. The occurrence of intermediates

between the subspecies makes the assignment of some

specimens difficult, but a combination of morphology

and geography is adequate for most material. Although

not perfect, this classification provides a basis from

which to work.

The type and other collections of Silene burchellii

from the Cape Peninsula and adjacent coastal regions

in the extreme southwestern parts of the Western Cape

have a distinctive facies that sets them apart, notably

the ± prostrate or decumbent stems and often broad

leaves, and we treat these plants as comprising a nar-

rowly circumscribed subsp. burchellii that is geographi-

cally isolated from other Western Cape populations by

the coastal Cape Fold Mountains. A second set of pop-

ulations from the coastal and near inland parts of the

northern Eastern Cape and KwaZulu-Natal and extend-

ing into Swaziland is distinctive in its broad leaves and

multiflowered inflorescences of nodding or half-pendent

flowers with very short calyces, mostly 10-12 mm long.

We treat these plants as subsp. multiflora. A third set of

populations with relatively long calyces, typically 20-25

mm long, is characteristic of the western, southern and

central parts of the subcontinent, occurring along the

western and southern Escarpment and the mountains of

the southwestern Western Cape, inland onto the interior

plateau. These populations are treated as subp. pilosel-

lifolia. The remaining southern African plants have mod-

erate-sized flowers, the calyx ( 12)15-18(-20) mm long,

and are treated as subsp. modesta. As accepted here,

this subspecies is widely distributed, from the Mpuma-

langa Escarpment southwards into the KwaZulu-Natal

Midlands and through the foothills of Eastern Cape and

KwaZulu-Natal Drakensberg, and northwards into tropi-

eal and North Africa.

Although Carl Thunberg seems to have been the first

to collect the species, during his visit to the Western

Cape in 1772-1775, he mistakenly associated his eol-

lection (UPS-THUNB 10780) with the European speeies

S. nutans L. (Table 1 ). Inexplicably, however, this name

was omitted from his Flora capensis (Thunberg 1794),

and the species was only recognized as distinct several

decades later (Otth 1824) from material collected on

Lion’s Head by William Burchell during his visit to the

region during 1810-1815.

Key to subspecies

la Stem.s prostrate or decumbent, extensively divaricately

branched with numerous axillary shoots, not strongly suf-

frutescent; leaves obovate-spathulate to oblanceolate, ±

densely puberulous; calyx 1 5-20 mm long; coastal regions

of the extreme southwestern Western Cape (? also Port

Elizabeth) 6a. subsp. burchellii

lb Stems suberect or shortly decumbent, brancing angle acute,

± sufifrutescent; leaves variable, upper usually narrower,

linear to narrowly oblanceolate, puberulous to subgla-

brous; calyx 10-25 mm long:

2a Leaves obovate to narrowly oblanceolate, lower 7-20 mm

wide; inflorescence mostly (5-)7-12-flowered, usually

with well-developed branch, lower intemodes 15-20 mm

long; flowers patent or slightly deflexed at anthesis; calyx

10-12(-15) mm long; coastal and near inland from E

Cape to S Mozambique and Swaziland ... 6d. subsp. multiflora

2b Leaves narrowly oblanceolate to linear-lanceolate or linear

(rarely lower leaves oblanceolate), mostly 2-5(-13) mm

wide; inflorescence mostly 5-8-flowered, lax or dense,

simple or branched, lower intemodes 15^0 mm long;

flowers mostly suberect, rarely patent; calyx (1 2-) 15-25

mm long, usually arcuate:

3a Calyx (18-)20-25 mm long; western and southwestern

coastal and interior southern Africa .... 6b. subsp. pilosellifolia

3b Calyx (12-)15-18(-20) mm long; southeastern, eastern and

northern southern Africa to N Africa and Arabia

6c. subsp. modesta

6a. subsp. burchellii

Silene tlmnbergiana Eckl. & Zeyh. ex Sond.: 128

(1860), syn. nov. Type: South Africa, [Western Cape],

‘steinige stellen der hohe am Tagelberge und Lowen-

berge’. Sept. [71832], Ecklon & Zeyher 253 (SAM,

lecto.!, designated by Boequet & Kiefer: 8 (1978); SAM

[2 sheets]!, SI, isolecto.).

Stems decumbent and extensively divaricately

branched with numerous axillary shoots, not strongly

sufifrutescent. Leaves obovate-spathulate to oblanceolate,

10-50 X (3-)6-22 mm, ± densely puberulous. Iriflores-

cence mostly 5-7-flowered, lax, lower intemodes 20-30

mm long. Flowers suberect. Calyx 15-20 mm long. Car-

pophore 6-10 mm long. Figure 13 A.

Distribution: coastal regions of the Western Cape

between Darling and Agulhas, and possibly at Port Eliz-

abeth in the Eastern Cape, occurring mainly on shale or

loamy soils in renosterveld but also in fynbos, on sand-

stone or limestone (Figure 14). On the Cape Peninsula,

subsp. burchellii is found only on Signal Hill and Table

Mountain, and is replaced by subsp. pilosellifolia on the

southern peninsula.

We include Paterson 2543 (PRE) from Port Eliza-

beth here although it represents the only collection of

subsp. burchellii from the Eastern Cape. The plants have

prostrate, well-branched stems, oblanceolate leaves,

and moderately sized flowers with calyx ± 1 5 mm long.

These features are anomalous for subsp. pilosellifolia,

the local form from the Port Elizabeth region.

Diagnosis: distinguished by the decumbent, divari-

cately branching stems with densely pubescent, obo-

vate to oblanceolate leaves rarely more than 4x as long

as wide; and moderately long calyx, 15-20 mm long.

Plants from Gansbaai and Agulhas are more compact

and densely leafy than usual, possibly in response to

their proximity to the ocean.

Silene ihunbergiana, based on plants collected essen-

tially in the same place as S. burchellii, was thought to

differ in its broader leaves but this apparent distinction

has no merit.

Bothalia 42,2(2012)

169

FIGURE 14. — Distribution of Silene biirchellii in southern Africa:

subsp. biirchellii. A; suhsp. pilosellifolia, o; subsp. modesta, •;

subsp. miiltiflora, A.

Dec. [without year], Ecklon & Zeyher 1959 (S, lecto.!,

designated here; PRE!, SAM!, isolecto.).

Silene hiirchellii var. cernua Rohrb.: 121 (1869),

nom. illegit. superfl. Type: as for S. biirchellii var.

angustifolia.

Silene dinteri Engl.: 383 (1912). Type: [Namibia],

‘auf der Granitkuppe bei Aus’, 11 Jan. 1910, Dinter

1141 (SAM, syn.l); 13 Apr. 1911, Dinter 2235 (SAM,

syn.i).

Silene cernua sensu Barth: 623 (1832), non Thunb.

(1794) (= S. aethiopica Burm.).

Silene cernua var. denudata Eenzl. in Drege: 222

(1844), nom. nud. [/zt/e Rohrb.: 121 (1869)]

Silene cernua var. hirta lus. linifolia Eenzl. in Drege:

222 (1844), nom. nud. [Specimen: South Africa, ‘ad

fl. Sunday’s River, pr. Nieweweld et in rupestribus as

Dutoitskloof, Drege 550a {fide Rohrb.: 121 (1869)]

Additional specimens

WESTERN CAPE. — 3318 (Cape Town): near Darling, (-AD),

Sept. 1905, Bolus 12615 (BOL); Tygerberg Nature Reserve, 200-400

m, (-DC), 13 July 1975, Loubser 3320 (NBG); ‘steinige lehmige stel-

len am Lbwenstaart [Lion’s Rump]’, (-DC). July without year, Eck-

lon & Zeyher 4748 (BOL, SAM); E slopes of Lion’s Head, (-DC),

Sept. 1913, Kensit s.n. (BOL); Signal Hill, (-DC), Sept. 1897, Thode

STE5830. 5831 (NBG); 28 Aug. 1938, Penfold 115 (NBG); Sig-

nal Hill, Schottsche Kloof, (-DC), Aug. 1938, Penfold 111 (NBG);

above Groote Schuur, (-DC), 12 Sept. 1895, Wolley Dod 93 (BOL);

path to Blockhouse, (-DC), 29 Sept. 1892, Guthrie 1182 (BOL);

Nursery Gorge, (-DC), 24 Oct. 1923, Compton s.n. (BOL); E slopes

of Table Mtn, (-DC), Sept. 1879, Bolus 4709 (BOL, PRE); sandy

places on Cape Flats, (-DC), without date or collector (NBG). 3418

(Simonstown): Constantiaberg, (-AA), 15 Dec. 1895, Wolley Dod 472

(BOL); N slope of Orange Kloof, (-AA), 6 June 1897, Wolley Dod

2616 (BOL); Hangklip, Skilpadsvlei, (-BD, 30' [9 m], 21 Oct. 1969,

Boucher 737 (NBG). 3419 (Caledon): Gansbaai, (-CB), Aug. 1940,

Stokoe 7600 (BOL). 3420 (Bredasdop); Cape Agulhas, (-CC), 27 Oct.

1940, Esterhuysen 4411 (BOL).

7EASTERN CAPE.— 3325 (Port Elizabeth): Port Elizabeth. (-

DC), Sept. 1914, Paterson 2543 (PRE).

6b. subsp. pilosellifolia (Cham. & Schltdl.)

J.C. Manning & Goldblatt, stat. nov. Silene pilosellifo-

lia Cham. & Schtdl. [as filosellaefolia'], Linnaea 1:

41 (1862). Silene burchellii var. pilosellifolia (Cham. &

Schltdl.) Sond. [as fiilosellaefolia'\. 128 (1860). Type:

South Africa, [Western Cape], ‘prope Plettenbergs-

bay ad Doukamma [Goukamma]’, Jul. 1821, Mund &

Maire s.n. (B, holo.t {fide Boucquet & Kiefer (1978)]).

Neotype: South Africa, Eastern Cape, Uitenhage Dist.,

sand-hills near the Zwartkopsrivier, Sept, [without year],

Zeyher 240 (NBG, neo.l, designated here; NBG!, SAM

[5 sheets]!, iso.). [The holotype appears to have been

destroyed (Bocquet & Kiefer 1978) and we accordingly

designate a neotype from the same geographical area as

the type and that matches both the protologue and the

current application of the name. Another collection that

has been identified as type material by the Stockholm

herbarium, Mund & Maire s.n. S-G-8699 (S), was col-

lected at Voormansbosch near Swellendam and thus has

no direct link to the protologue at all.]

Silene burchellii var. angustifolia Sond.: 128 (1860).

Type: South Africa, [Western Cape], Caledon Zwartberg,

Stems ± erect or shortly decumbent, ± tufted with

branches subbasal and at an acute angle, ± suffrutes-

cent. Leaves oblanceolate to linear-lanceolate or linear

but lower leaves sometimes broader, 20-55 x 2-5(-13)

mm, puberulous or subglabrous. Inflorescence mostly

3-8-flowered, lax or dense, lower intemodes 15^0 mm

long. Flowers mostly suberect, rarely patent. Calyx ( 1 8-

)20~25 mm long, usually flexed slightly upwards. Car-

pophore 9-1 1 mm long, figure 13B.

Distribution and ecology’: widely distributed through

western, southern and the eastern interior of south-

ern Africa, from central Namibia southwards along the

western escarpment, through the Cape Fold Mountains

into Eastern Cape as far as Zuurberg and inland onto the

eastern interior plateau through Free State and Gauteng

as far north as Limpopo (Figure 14), occurring in a wide

range of open shrubby or grassy habitats, mostly among

rocks.

Diagnosis: distinguished from other subspecies by its

± suberect, mostly longer flowers, the calyx ( 1 8-)20-25

mm long and curved slightly upwards at the junction

between anthophore and ovary. This slight curvature

is lost in fruit. Plants are suffrutescent and variable in

foliage, usually with oblanceolate to linear-lanceolate

or linear leaves although the lower leaves are some-

times broader. Populations from the western edge of the

Escarpment in Namibia and Northern Cape southwards

into the Cedarberg in Western Cape have sub-glabrous

leaves.

The distinction between subsp. pilosellifolia, with

calyx ( 1 8-)20-25 mm long and subsp. modesta, with

calyx (12-)15-18(-20) mm long is not always a very

easy one to draw and even somewhat arbitrary in some

cases.

Two collections from the mountains above Simons-

town in the southern Cape Peninsula {Adamson 2333

& Esterhuysen 34106) are included here on account

of their suberect stems and caespitose habit, narrowly

oblanceolate leaves, and long calyx, ± 22 mm long.

They appear to represent isolated outliers and are the

170

Bothalia 42,2 (2012)

only records of subsp. pilosellifolia from the Penin-

sula— other populations from north of Constantiaberg

represent subsp. burchelUi.

Populations from the summit of Mariepskop [Van der

Schijff 4882 (PRE), Van der Schijff 5594 (PRE), Hardy

7000 (PRE)] and from the Wolkberg along the north-

eastern escarpment {Miiller & Scheepers 128 (PRE)]

appear to represent a distinctive ecotype with prostrate

or sprawling, highly branching stems, forming tangled

mounds, and broad, spathulate leaves. The Mariepskop

population was to have formed the basis of the unpub-

lished manuscript name S. junodii D. Masson ms. At this

stage, however, it is unclear how much of this morpho-

logical differentiation is phenotypic since these plants

invariably occupy sheltered, mostly shaded situations

among rocks, where they avoid fires. They are provi-

sionally included in subsp. pilosellifolia on account of

their calyx length, 20-23 mm long.

Additional specimens

NAMIBIA. — 2014 (Welwitschia): Spitskop, (-BA), 25 Sept. 1981,

Muller 1601 (WIND). 2217 (Windhoek): Grossherzog Friedrichsberg,

Farm Regenstein, (-CA), 2 336 m, 19 Mar. 1972, Giess 11681 (NBG,

PRE, WIND); 18 km from Windhoek, (-CB), 27 May 2000, Zimmer-

mann 239 (WIND). 2616 (Aus): Aus, mountain on Farm Klein Aus,

(-CA), 18 Aug. 1963, Merxmuller 2956 (WIND); Aus, (-CB), 9 Jul.

1922, Dinter 3579 (BOL, PRE, WIND); 4 Mar. 1929, Dinter 6109

(BOL, NBG); on road P705, 26 Oct. 1987, Kolbei-g 240 (WIND).

2715 (Bogenfels): NW of beacon, (-BD), 9 Sept. 1992, Oliver 10198

(WIND). 2716 (Bethanie); Karas, E of summit, (-CD), 21 Sept. 1977,

Merxmuller 32193 (WIND); slope of mountain, (-DD), 24 Sept. 1972,

Merxmuller 28811 (WIND). 2718 (Griinau): Klein Karas, (-CA), 9

Apr. 1931, Ortendahl 50 (PRE, WIND).

LIMPOPO. — 2328 (Baltimore): Blouberg, (-BB), 1 June 1953,

Esterhuysen 21437 (BOL); 20 Feb. 1990, Stirton, Venter & Edwards

12680 (NU). 2430 (Pilgrim’s Rest): Wolkberg, Serala Peak, (-AA), 23

Apr. 1971, Miiller & Scheepers 128 (PRE).

GAUTENG. — 2528 (Pretoria): top of hill behind Louis Botha’s

home, (-CA), 12 Oct. 1925, Smuts 870 (PRE); 9 miles [14.4 km], E of

Pretoria, The Willows Farm, (-CA), 10 Nov. 1946, Codd 2139 (PRE);

Willowglen, (-CA), Jan. 1951, Forssman 6 (PRE); Botanical Research

Institute, (-CB), 10 Dec. 1975, Scott 3 (PRE). 2628 (Johannesburg):

Doomkloof, (-CC), 25 Nov. 1928, Gillett 2555 (BOL); 3 Dec. 1928,

Gillett 207 (NBG); 28 Dec. 1928, Gillett 3246 (BOL).

MPUMALANGA. — 2430 (Pilgrim’s Rest): Mariepskop summit,

(-DB), 5 Jan. 1960, Van der Schijff 4882 (PRE); 6 Jul. 1961, Van der

Schijff 5594 (PRE); 3 Sept. 2000, Burrows 7013 (NU); Mariepskop

Radar Station, (-DB), 16 Feb. 1990, Hardy 7000 (PRE). 2529 (Wit-

bank): Middelberg Dist., Buffelsvlei, (-BC), I Dec. 1933, Rudatis 6

(NBG); Middelburg, (-CD), without date, Guthrie 4064 (NBG).

FREE STATE. — 2627 (Potchefstroom): II miles [17.6 km] from

Parys to Potchefstroom, (-DD), 8 Apr. 1967, Vahrmeyer 1568 (PRE).

2827 (Senekal): Ficksburg, (-DD), 26 Oct. 1934^ Galpin 13874

(BOL); Strathcona, (-DD), 7 Nov. 1936, Fawkes 12 (NBG). 2926

(Bloemfontein): Dewetsdorp, (-DA), 15 Apr. 1950, Steyn 904, 945

(NBG). 3027 (Lady Grey): Zastron Dist., (-AC), Nov. 1926, Maree 57

(PRE).

KWAZULU-NATAL.— 2829 (Harrismith): foot of Griffin’s Hill, (-

DD), 2 Apr. 1945, Acocks 11387 (PRE). 2830 (Dundee): Mpate Mt.,

(-AA), 26 Nov. 1964, Shirley s.n. (NU).

LESOTHO. — 2828 (Bethlehem): Leribe, (-CC), without date,

Dieterlen 362 (PRE, SAM); Dieterlen 610 (PRE).

NORTHERN CAPE. — 2816 (Oranjemund): Richtersveld, Rem-

hoogte, (-BD), 13 Sept. 1929, Herre 11778 (NBG); Numees Camping

Site, (-BD), 26 Sept. 1981, Hugo 2787 (NBG, PRE); W side of ridge

N of Numees Camp, (-BD), 19 Sept. 1981, McDonald 688 (NBG);

Numees Mountain, (-DB), Sept. 1995, Williamson <& Williamson 5816

(NBG). 2817 (Vioolsdrif): Koeboes [Khubus], (-AC), 17 Sept. 1929,

Herre H779 (NBG); Rosyntjieberg, neck N of Lclieshoek, (-AC),

1 060 m, 30 Aug. 1977, Oliver, Tolken & Venter 300 (NBG, PRE);

Koeskop, (-AC), Sept. 1995, Williamson & Williamson 5779 (NBG);

Zebrakloof, NE of Rosyntjieberg, (-AC), 9 Oct. 1991, Germishuizen

5560 (PRE); Ploegberg S of Khubus, (-CA), 600 m, 3 Sept. 1977,

Oliver, Tolken & Venter 509 (NBG, PRE); Ploegberg, (-CA), 25 Sept.

1991, Van Jaarsveld 11931 (PRE). 2917 (Springbok): Spektakel, (-

DA), 26 Aug. 1941, Compton 11534 (NBG). 2918 (Gamoep): 15 miles

[24 km] NE of Springbok, (-AA), 8 Sept. 1950, Maguire 351 (NBG).

3017 (Hondeklipbaai): Farm 477 Taaibosduin, 20 Aug. 2009, Bester

9611 (NBG, PRE); Grootvlei, (— BB), 7 Sept. 1945, Compton 17280

(BOL, NBG); Skilpad Flower Reserve, (-BB), 3 Oct. 1995, Cruz 132

(NBG). 3018 (Kamiesberg): Langkloof, N of Farm Doringkraal, (-

CA), 23 Sept. 2010, Goldblatt & Porter 13574 (NBG). 3120 (Willis-

ton): N of Farm De Hoop, (-CC), 7 Jan 1986, Snijman 991 (NBG).

WESTERN CAPE. — 3118 (Vanrhynsdorp): Knersvlakte, ± 30 km

N of Vanrhynsdorp, Farm Ratelgat, limestone ridge ± 400 m SE of

Matjieshuise, (-BC), 7 Aug. 2011, Koopman s.n. (NBG). 3119 (Cal-

vinia): upper part of Vanrhyn’s Pass, (-AC), 800 m, 13 Sept. 1993,

Strid & Strid 37844 (NBG); parking near top of Vanrhyn’s Pass, (-AC),

13 Sept. 2010, Goldblatt & Porter 13527 (MO, NBG). 3218 (Clanwil-

liam): Piketberg, Zebra Kop, (-DB), 16 Dec. 1979, Esterhuysen 35331

(BOL). 3219 (Wuppertal): Kliphuis Gully leading up to Pakhuis Peak,

(-AA), 22 Oct. 1987, Taylor 11863 (NBG); Sneeugat, (-AA), 15 Jan.

1923, Andreae 922 (PRE); Elandskloof, (-CA), 30 Sept. 1936, Leipoldt

s.n. BOL31661 (BOL); Middelberg Plateau, (-CA), 14 Dec. 1941, Bond

1351 (NBG). 3318 (Cape Town): Jonkershoek, Guardian Peak, (-DD),

[month illegible] 1946, Adamson 3654 (BOL); Jonkershoek, (-DD),

18 Sept. 1936, Borchardt 440 (PRE). 3319 (Worcester): Groot Win-

terhoek, (-AA), 14 Feb. 1934, Compton 4628 (BOL); Mostertshoek

Twins, (-AD), Jan. 1944, Esterhuysen 9842 (BOL); Hex River Mtns,

Sentinel Peak, (-AD), 16 Feb. 1958, Esterhuysen 27572 (BOL); Groot

Drakenstein Mtns, Duiwelskloof, (-CC), 12 Dec. 1943, Wasserfall 742

(NBG); Haelhoeksneukop, (-CC), 16 Dec. 1975, Esterhuysen 34161

(BOL). 3320 (Montagu): Laingsburg Dist., Whitehill Ridge, (-BA),

10 Nov. 1935, Compton 5900 (BOL, NBG); Cabidu, (-BB), 28 Oct.

1950, Compton 22213 (NBG); Grootvadersbos State Forest, Boos-

mansbos Wilderness Area, (-DD), 30 Nov. 1988, Van der Merwe 284

(PRE). 3321 (Ladismith): Gamka Mtn Reserve, (-BC), 25 Nov. 1975,

Boshoff P275 (NBG); Touwsberg, Farm Rietfontein, (-CA), 7 Oct.

1993, Smook 8705 (PRE). 3322 (Oudtshoom): Swartberg Pass, (-AC),

7 Aug. 1949, Steyn 267 (NBG); 6 Dec. 1987, Vlok 1895 (PRE); near

top of Swartberg Pass, (-AC), 21 Mar. 1976, Thompson 2762 (NBG,

PRE); upper Cango valley, Bassonsrus, (-AC), 4 November 1974,

Moffett 433 (NBG); Meiringspoort, (-BC), 12 Nov. 1941, Thorne s.n.

(NBG). 3418 (Simonstown): hills above Simonstown, edge of swamp,

(-AA), 11 Dec. 1938, Adamson 2333 (BOL); Swartkops Peak S of

Simonstown, on firebelt above Froggy Pond, (-AA), 16 Nov. 1975,

Esterhuysen 34106 (BOL); Harold Porter Botanic Garden, above Disa

Kloof, (-BD), 17 Oct. 1991, Forrester 974 (NBG). 3419 (Caledon):

Caledon Baths, (-AB), July 1892, Guthrie 2469 (NBG); between Cal-

edon and Napier, (-BD), 3 Aug. 1940, Esterhuysen 3051 (BOL); ±12

km NW of Napier, Fairfield Farm, (-BD), 6 Oct. 1994, Kemper 1PC632

(NBG). 3420 (Bredasdorp): Adamskop, (-AC), 200 m, 15 Oct. 1982,

Bayer 31 99 Q4BG).

EASTERN CAPE. — 3125 (Steynsburg): Middelburg Dist., Bangor

Farm, (-AC), Oct. 1917, Bolus 14040 (BOL). 3126 (Queenstown):

Broughton near Molteno, (-AD), Dec. 1892, Flanagan 1567 (NBG,

PRE, SAM); Queenstown, (-DD), Feb. 1896, Galpin 1988 (PRE); Jan.

1 962, Bokelmann 1 (NBG); 1 5 km from Dordrecht on road to Queens-

town, (-DB), 13 Jan. 1997, Germishuizen 8891 (PRE). 3323 (Willow-

more): between Willowmore and Patensie, Nuweveld Pass, (-BC), 1 5

Sept. 1982, Balkwill 451 (NU); Langkloof, near Misgund, (-DC), Oct.

1921, Fourcade 1718 (BOL). 3224 (Graaf-Reinet): Schimper’s Hill

near Graaf-Reinet, (-BA), 4 July 1865, without collector BOL74032

(BOL). 3324 (Steytlerville): 51.1 miles [82 km] from Humansdorp on

road to Willowmore, (-CA), Oct. 1934, Fourcade 5169 (NBG); Zuur

Anys Hills [Suuranysberge], (-CC), Dec 1932, Fourcade 4945 (BOL);

kloof on road to Hankey, (-DD), 12 Sept. 1942, Fourcade 5746 (BOL,

NBG). 3325 (Port Elizabeth): Zuurberg, (-BD), 27 May 1965, Bayliss

2686 (NBG); 8 Jan. 1986, Van Wyk & Van Wyk 1183 (PRE).

6c. subsp. modesta J.C. Manning & Goldblatt, subsp.

nov.

TYPE. — Eastern Cape, 3227 (Stutterheim): Mount

Kemp, (-CB), 13 Jan. 1947, Compton 19196 (NBG,

holo.).

Bothalia42,2 (2012)

171

Silene burchellii var. latifolia Sond.: 128 (1860), syn

nov. Type: South Africa, [Gauteng], shady places at the

Crocodile River, Zeyher s.n. S] 1-24323 (S, holo.!).

Silene acuta E.Mey. in Drege: 222 (1844), nom.

nud. [Specimen: South Africa, [Eastern Cape], between

Omtendo [Mtentu] and Omsamculo [uMzimkhulu],

Drege 5340 [fide Sond.: 128 (I860)].

Silene cernua var. hirta lus. lanceolata Fenzl. in

Drege: 222 (1844), nom. nud. [Specimen: South Africa,

[Eastern Cape], ‘iter Gekan et Busche’, Drege 550h [fide

Rohrb.: 121 (1869)].

[For synonyms from tropical and North Africa see

Turrill (1956a), Wild (1961), Wickens (1976) and Gil-

bert (2000)].

Stems ± erect or very shortly decumbent, ± tufted

with branches subbasal and at an acute angle, ± suf-

frutescent. Leaves oblanceolate to linear-lanceolate

or linear but lower leaves sometimes broader, 20-55 x

2-5(-13) mm, puberulous or subglabrous. Inflorescence

mostly 3-8-flowered, lax or dense, lower intemodes

15^0 mm long. Flowers mostly suberect, rarely patent.

Calyx (12-)15-18(-20) mm long, usually flexed slightly

upwards. Carpophore 6-9(-10) mm long.

Distribution and ecology, widely distributed through

eastern southern Africa, Lesotho and Swaziland north-

wards into tropical and North Africa and Arabia (Figure

14), essentially in temperate or subtropical grassland and

savanna, often in rock outcrops.

Diagnosis: subsp. modesta constitutes the core of this

variable species after the more extreme forms have been

segregated and includes ± suffhitescent, tufted plants

with mostly oblanceolate to linear leaves and suberect,

moderately-sized flowers with calyx (12-)15-18(-20)

mm long. This taxon seems to have included the bulk

of the collections that were to have been recognized as

a new species under the unpublished manuscript name

Silene australis D. Masson ms.

Subsp. modesta is replaced in southern Mozambique

and coastal KwaZulu-Natal and Eastern Cape by subsp.

multiflorus, mostly with broader leaves and with gener-

ally shorter flowers, the calyx mostly 12-15 mm long,

and in the western, southern and eastern interior of the

subregion by subsp. pilosellifolia with mostly longer

flowers, the calyx 18-25 mm long. Subsp. burchellii,

with procumbent or decumbent stems, is restricted to the

extreme southwestern coastal regions.

The tropical African material, all with a calyx length

of 12-20 mm, is comfortably included in subsp. mod-

esta. Previously aberrant populations from northern

Kenya, eastern Uganda, and Ethiopia with exception-

ally long calyces, 26-35 mm long, treated as var. gillettii

Turrill (1954, 1956a), have since been segregated as the

distinct species Silene gillettii (Turrill) M.G.Gilbert, also

distinguished by its broader leaves with broadly cuneate

or subcordate bases (Gilbert 2000).

Additional southern African specimens

LIMPOPO. — 2329 (Pietersburg): Farm Duvenhageskraal, (-CD/

DC), 3 Dec. 1985, Venter 11283 (NU). 2330 (Tzaneen): Woodbush

Forestry Station, (-CC), Jan. 1923, Wager 23094 (PRE); 23 Dec.

1928, Gillett 3205 (NBG). 2428 (Nylstroom): Geelhoutkop, (-AD),

Jan. 1918, Breijer 18089 (PRE); Warmbaths, (-CD), 30 Sept. 1908,

Leendertz 1352 (PRE).

NORTH-WEST. — 2526 (Zeerust): Lichtenburg Dist., Grasfon-

tein, (-CC), 16 Dec. 1929, Sutton 343 (PRE). 2527 (Rustenburg):

Rustenburg Dist., (-BA), without date, Mudd s.n. (BOL); Rustenburg

Nature Reserve, (-CA), 8 Oct. 1970, Jacobsen 1067 (PRE). 2626

(Klerksdorp): Ventersdorp, (-BD, Feb. 1932, Wilman NBG252/30

(BOL).

GAUTENG. — 2528 (Pretoria): Fountains Valley, (-CA), 14 Nov.

1928, Repton 118 (PRE); Wonderboom Reserve, (-CA), 20 Oct. 1944,

Repton 1887 (PRE); 14 miles [22.4 km] SE of Pretoria on road to Del-

mas, (-CA), 21 Dec. 1950, Codd 6302 (PRE). 2627 (Potchefstroom):

Northcliffe, (BB), 26 Oct. 1984, Behr 785 (PRE). 2628 (Johan-

nesburg): ridge above Jeppestown, (-AA), Feb./Mar. 1894, Galpin

1376 (PRE); Kensington Ridge, (-AA), 31 Oct. 1932, Burn Davy 57

(NBG); Melville Koppies, (-AA), 3 Oct. 1960, Macnae 1265 (BOL);

Mondeor, (-AA), 9 Nov. 1961, Lucas 35146 (PRE); Aasvogels Kop,

(-AA), 24 Apr. 1927, Young 26444 (PRE).

MPUMALANGA. — 2430 (Pilgrim’s Rest): Lekgalameetse Nature

Reserve, The Downs, (-AA), 21 Jan. 1986, Stalmans 1005 (PRE); 19

Dec. 1988, Stahlmans 1811 (PRE); Mt Saheba Nature reserve, (-DC),

Jan. 1976, Forrester & Gooyer 46 (PRE); Graskop, (-DD), 3 Dec.

1887, Galpin s.n. (BOL); Stanley Bush Kop, (-DD), 9 Dec. 1986,

Raal & Raal 1016 (PRE). 2530 (Lydenburg): Dullstroom, (-AC), 30

Jan. 1959, Werderman & Oberdieck 2045 (PRE); 24 Feb. 1982, Cam-

eron 151 (PRE); Mt. Anderson, (-BA), 25 Dec. 1932, Smuts & Gillett

2468 (BOL, PRE); Farm de Kuilen, (-BA), 13 Mar. 1985, Krynaim

313 (PRE); Mac Mac Nature Reserve, (-BB), 7 Mar. 1979, kluge 1789

(PRE); Belfast, (-CA), 31 Jan. 1929, Hutchinson 2765 (BOL); 16 Feb.

1964, Bayliss 2022 (NBG); Dullstroom Dist., Farm Macduff, (-CB),

24 Feb. 1989, Burgoyne 1053 (PRE); Farm Uitkomst, 18 km from

Machadadorp on road to Badplaas, (-CD), 5 Mar. 1986, Germishuizen

3811 (PRE); Kaapsehoop Asbestos Mine, (-DB), 1 Mar. 1987, Morrey

& Cadman 3679 (NU). 2531 (Komatipoort): Barberton Dist., Lomati

Valley, (-CC), Feb. 1922, Thorncroft 1123 (PRE); summit of Saddle-

back Mtn, Farm Dycedale, (-CC), 18 Apr. 1987, Brusse 5053 (PRE).

2630 (Carolina): Carolina Dist., (-AA), 18 Nov. 1909, Rademacher

8195 (PRE); Ermelo Dist., Farm Nooitgedacht, (-AC), 9 Mar. 1937,

Henrici 1601 (PRE); Songimvelo Game Reserve, between Hoog-

genoeg and Lochiel, (-BB), 11 Dec. 1992, Jordaan 2520 (PRE). 2729

(Bethal): Amersfoort, Welverdacht Farm, (-DB), 15 Mar. 1985, Turner

692 (PRE); Ermelo, (-DB), 1906, Burtt Da\y. 5453 (PRE); Feb. 1910,

Leendertz 7807 (PRE). 2730 (Vryheid): E of Wakkerstroom, (-AD),

13 Jan. 1986, Glen 1589 (PRE); near Piet Retief, (-BB), 13 Jan. 1951,

Compton 22339 (NBG).

FREE STATE. — 2828 (Bethlehem): Clarens, (-CB), Nov. 1917,

Van Hoepen 18185 (PRE); Golden Gate, (-DA), 15 Jan. 1976, Fenn

6 (NU); Golden Gate National Park, Mt Pierre, (-DA), 8 Jan. 1989,

Groenewald 8711 (PRE); Golden Gate Highland Park, (-DA), Jan.

1966, Liebenberg 7495 A (PRE); Witzieshoek, (-DB), Dec. 1905,

Thode STE5663 (NBG). 2829 (Harrismith): Van Reenen, (-AB), Jan.

1914, Bews 962 (NU); Harrismith, Queen’s Hill, (-AC), 22 Feb. 1969,

Jacobsz 1073 (PRE); Harrismith, Botanic Garden, 5 500' [1 667 m],

(-AC), 18 Jan. 1970, Van der Zeyde s.n. (NBG); Drakensberg Botanic

Garden, (-AC), 23 Jan. 1975, Jacobsz 2093 (NBG, PRE); Swinburne,

Farm Grootvlei, (-AC), 30 Jan. 1961, Jacobsz 9 (PRE); Sterkfontein

Dam, (-AC), 28 Nov. 1974, Jacobsz 1858 (PRE); 11 Dec. 1985, Blom

333 (PRE). 2926 (Bloemfontein): Wintervalley, N of Bloemfontein, (-

AA), 6 Dec. 1968, Muller 386 (PRE); Bloemfontein, O.F.S. Botanic

Garden, (-AA), 6 Dec. 1968, Muller 386 (NBG); Grant’s Hill near

Oranje School, (-AA), 23 Oct. 1925, Potts 3526 (PRE).

SWAZILAND. — 2631 (Mbabane): hilltop near Mbabane, (-AC),

20 Feb. \96\, Dlamini s.n. (NBG, PRE).

KWAZULU-NATAL.— 2729 (Volksrust): Newscastle Dist., Blue

Ridge, (-DD), 25 Nov. 1989, Smit 1315 (PRE). 2730 (Vryheid):

Hlobane, (-DB), 5 Mar. 1950, Johnstone 389 (NU). 2731 (Louws-

burg): Itala Game Reserve, (-CB), 17 Jan. 1978, McDonald 475 (NU);

19 Oct. 1982, Germishuizen 2235 (PRE); Ngome, (-CD), 3 Apr. 1977,

Hilliard & Burtt 9961 (NU); Ngome, Ntendeka Wilderness area, (-

CD), 4 dec. 1985, Jordaan 580 (PRE). 2828 (Bethlehem): Royal Natal

National Park, 200 m along path from Basuto Gate to Tendele, (-DB),

5 Feb. 1982, Manning 87 (NU); Mont aux Sources, (-DD), Apr. 1920,

Allsop 34 (PRE). 2829 (Harrismith): Little Switzerland, (-CB), 21 Apr.

1969, Anderson 227 (PRE). 2929 (Underburg): Giant’s Castle Game

172

Bothalia 42,2 (2012)

Reserve, (-AD), 26 Jan. 1966, Traiiseld 542 (NU); Mooi River, (-BB),

Dec. 1942, Fisher 419 (NU); Mooi River Dist., Hidcote, (-BB), 15

Jan. 2003, Potgieter 876 (NU); Kamberg, (-BC), 31 Dec. 1974, Wright

2090 (NU); Mpendle Dist., Mulangane Ridge above Carter’s Nek, (-

BC), 3 Feb. 1984, Hilliard & Burt! 17 523 A (NU, PRE); Loteni Nature

Reserve, (-BC), 13 Dec. 1978, Phelan 223 (NU); Sani Pass, (-CB),

21 Mar. 1977, Hilliard & Burtt 9743 (NU); headwaters of Mhlahl-

angubo River, (-CB), 23 Jan. 1982, Hilliard & Burtt 15343 (E, NU);

Bulwer Dist., Mahwaqa Mt., (-DC), 11 Jan. 1992, Feltham 124 (NU);

Polela Dist., Farm Sunset, (-DC), 30 Dec. 1973, Rennie 445 (NU); 8

Feb. 1985, Rennie 1616 (NU). 2930 (Pietermaritzburg): Karkloof, Mt.

Gilboa, (-AD), 17 Dec. 2000, Johnson & Neal 33 (NU); Noodsberg,

Lager Farm, (-BD), 14 Oct. 1989, Williams 558 (PRE); Table Mt.,

(-CB), 7 Jan. 1949, Killick 220 (NU); Byrne Valley, (-CC), 31 Oct.

2002, Potgieter 841 (NU); Hella Hella, 3 800' [1 160 m], (-CC), 1

Nov. 1997, Ingomankulu Hill, (-CD), 15 Dec. 2004, Young 208 (NU).

LESOTHO. — 2927 (Maseru): Molimo-Nthuse on way to Pass,

(-BC), 20 Oct. 1975, Schmitz 6239 (PRE). 2828 (Bethlehem): Butha

Buthe, (-CC), 9 500' [2 878 m], 2 Feb. 1954, Coetzee 409 (NBG,

PRE); Malibamatzo Valley, (-DC), Dec. 1971, Schmitz 1545 (PRE);

New Oxbow Lodge, (-DC), 13 Jan. 2003, bester 3937 (PRE). 2928

(Marakabei): from Ha Lephoi Village along Lesobeng River, (-CB),

5 Mar. 1990, Smook 7271 (PRE), 2929 (Underberg): Sehlabathebe

National Park, (-CC), 7 Jan. 1979, Hoener 2121 (PRE).

WESTERN CAPE. — 3221 (Merwewille): Roggeveld, Fanri

Uitkyk, (-AD), Oct. 1920, Marloth 9704 (PRE); Beaufort West Dist.,

Layton, (-BB), 24 Nov. 1983, Shearing 400 (PRE). 3222 (Beaufort

West): Beaufort West Dist., Nieuweveld Mtns, (-BD), July 1895, Mar-

loth 2136 (PRE); Nieuweveld Mtns, Mountain View Farm, (-BD), 18

Apr. 1978, Gibbs Russel, Robinson & Herman 482C (PRE).

EASTERN CAPE.— 3027 (Lady Grey): Witteberg, (-CB), 11

Mar. 1904, Galpin 6583 (PRE); Rhodes, Carlisle’s Hoek, (-DB), 27

Dec. 1977, Bigalke 22 (NU). 3127 (Lady Frere): 25 km from Cala at

turnoff to Engcobo, (-DB), 12 Jan. 1997, Germishuizen 8819 (PRE).

3128 (Umtata): Ugie Dist., Pomona, (-AA), Feb. 1928, Gill 178

(NBG); Elandslaagte, (-AA), Mar. 2000, Edwards & Potgieter 1922

(NU); Umtata, (-DB), 18 Jan. 1895, Schlechter 6326 (PRE). 3226

(Fort Beaufort): top of Katberg Pass at Devil’s Bellows, (-BC), 24

Apr. 1995, Victor 1183 (PRE); ± 5 km above Nico Malan Pass, (-BD),

7 Feb. 1995, Victor & Hoare 324 (PRE). 3227 (Stutterheim): Thomas

River, (-AD), 15 Jan. 1947, Compton 19309 (NBG); Leighton 2797

(BOL); King William’s Town, (-CB), Nov. 1892, Sim 966. 967 (NU);

East London Dist., Amalinda Nature Reserve, (-DD), 22 Oct. 1984,

Currie 9 (PRE). 3326 (Grahamstown): Grahamstown, (-BC), Oct.

1888, Galpin 224 (PRE); Martindale, (-BD), 30 Nov. 1955, Tavlor

4962 (NBG); 7 Jan. 1965, Bayliss 2600 (NBG).

6d. subsp. multiflora J.C. Manning & Goldblatt,

subsp. nov.

TYPE. — KwaZulu-Natal, 2930 (Pietermaritzburg):

Krantzkloof, (-DD), Nov. 1921, Haygarth 9437 (NBG,

holo.; NH, PRE, iso.).

Stems ± erect or very shortly decumbent, ± tufted

with branches sub-basal and at an acute angle, ± suf-

frutescent. Leaves obovate to narrowly oblanceolate,

20-55 X (2-)7-15(-20) mm, upper leaves usually nar-

rower, puberulous or subglabrous adaxially. Inflores-

cence mostly (5-)7-12-flowered, dense, lower inter-

nodes 15-20 mm long (exceptionally lowermost 40 mm

long). Flowers nodding or slightly deflexed at anthesis.

Calyx 10-12(-15) mm long. Carpophore 3-6 mm long.

Distrihutioir. coastal and near inland from the Fish

River Mouth in Eastern Cape northwards through Kwa-

Zulu-Natal into southern Mozambique and inland to

near Mbabane in Swaziland, occurring mainly in sub-

tropical grasslands on loamy or sandy soils (Figure 14).

Diagnosis: distinguished by the shortly decumbent

or ± tufted habit with relatively broad, oblanceolate to

obovate lower leaves, and the dense, multi-flowered

inflorescence (typically with a well-developed branch),

of small, nodding or slightly pendent flowers with short

calyces, 10-12(-15) mm long.

The distinction between subsp. multiflora and subsp.

modes ta is blurred in the KwaZulu-Natal midlands,

where the transition between subsp. multiflora with

broad-leaves and subsp. modesta with narrowly oblan-

ceolate leaves, mostly less than 5 mm wide, lies along

the Drakensberg foothills at an altitude of ± 1 500 m.

Coastal forms of subsp. multiflora with more decum-

bent branches may be difficult to separate from Silene

crassifolia subsp. primuliflora in southern KwaZulu-

Natal and Eastern Cape where the distribution of the two

taxa overlaps. Introgression between the two species is

one possible cause. An alternative hypothesis is that S.

crassifolia is a coastal dune derivative of S. burchellii,

with the two forms becoming progressively more dis-

tinct to the south as ecological and geographical isola-

tion between them increases. Plants with the following

combination of characters are treated as subsp. multi-

flora: stems not strongly prostrate, branching angles

acute; inflorescence branched and multi-flowered with

more than five flowers, well separated from foliage by

leafless stem ±100-150 mm long bearing one or two

pairs of bract-leaves; calyx 10-12(-15 mm) long; cap-

sules ovoid, 8-10 X 5-7 mm.

South African material of this taxon has tradition-

ally been identified as var. latifolia Sond. but this name

is typified by a specimen from Gauteng and correctly

applies to broad-leaved forms of subsp. pilosellifolia.

It was to have been recognized at species level by Da-

niel Masson, and several herbarium specimens bear the

unpublished name Silene natalensis D. Masson ms.

Additional specimens

SWAZILAND.— ^2631 (Mbabane): Mbabane, (-AC), 20 Dec.

1952, Compton 23795 (NBG); Ukutula, (-AC), 29 Oct. 1954, Comp-

ton 24597 (NBG); 13 Nov. 1955, Compton 25249 (NBG).

MOZAMIBIQUE. — 2632 (Bela Vista): road from Ponta Molan-

gane to Ponta Do Puro, (-BC), 25 Nov. 2001, Govender 72 (NH);

Maputo Elephant Park, (-BC), 29 Nov. 2001, Govender 90 (NH).

KWAZULU-NATAL.— 2632 (Bela Vista): Kosi Bay, (-DD), 28

Sept. 1961, Meyer s.n. (NU); road from Kosi Estuary to Kosi Lake

Campsite, 50 m, (-DD), 18 Nov. 1982, Balkwill 571 (NH, NU); Kosi

Bay, Natal Parks Board Camp, (-DD), 27 Nov. 1967, Strey & MoU

3939 (PRE); Kosi Mouth, (-DD), 24 Apr. 1995, Lubbe 646 (NU).

2731 (Louwsburg): Nhloenkulu Mission Station, (-DC), July 1927,

Markotter s.n. STE8682 (NBG). 2732 (Ubombo): NW of Sibaya, (-

BC), 21 Sept. 1995, Lubbe 717 (NU); Lake Sibaya, (-BC), 16 Sept.

1965, Vahrmeijer 1122 (PRE); Mazengwenya, ± 6 km W of Mabibi,

(-BC), 17 Sept. 1994, Lubbe 273 (NH); Mazengwenya, E of Vazi

Swamp, (-BC), 27 Nov. 1969, MoU 4725 (NH); Mbazwane/Sodwana

Rd., (-DA), 28 Nov. 1971, Pooley 1516 (NU); Mbazwana, (-DA), 27

Sept. 1977, Balsinhas 3270 (PRE); St. Lucia Bay, (-DC), 26 Jul. 1939,

Schweickerdt 1371 (NH, PRE); I km N of Lake St. Lucia, (-DC),

24 Sept. 1987, MacDevetle 1892 (NH). 2831 (Nkandla): Melmoth,

Mooiplaas, (-CA), ± 930 m, 10 Oct. 1995, Hutchings <& Williams 3554

(NH). 2832 (Mtubatuba): Hluhluwe Game Reserve, (-AA), 5 Oct.

1953, Ward 1519 (NU); W of Charter’s Creek, (-AC), 5 Dec. 1955,

Ward 2840 (NH, NU, PRE); Hlabisa, Mfolozi flood plain, (-AC), 20

oct. 1985, Steyn 1 1 (NH); Dukuduku, (-AC), 25 Nov. 1965, Strey

6098 (NH, NU, PRE); 24 Jul. 1986, Steyn 55 (NH); Lake St Lucia,

East Shores, (-BA), 11 Oct. 1974, Taylor 238 (NU); Richard’s Bay,

(-BD), 14 Jul. 1929, Rump s.n. (NH, NU); 6 July 1974, Ward 8670

(NU, PRE); 16 Jul. 1974, Ward 2490 (NU). 2930 (Pietermaritzburg):

Greytown, Blinkwater Trails area, (-BA), 1 dec. 2001, Potgieter 560

(NU); Pietemiaritzburg, (-CB), Nov. 1939, Thomas 14 (NBG); Pie-

Bothalia 42,2 (2012)

173

termaritzburg. Town Hill, (-CB), Oct. 1942, Fisher 362 (NU); Nov.

1944, Fisher 741 (NH); 30 Oct. 1952, Compton 23 738 (NBG); Thom-

ville, (-CB), 7 Nov. 1964, Shirley s.n. (NU); Drummond, (-CC), Oct.

1929, Rump s.n. (NU); Mid lllovo, (-CC), 10 Dec, 2008, Young 863

(NU); Inanda, (-DA), Nov. 1884, Medley Wood 393 (NBG, NH); Aug.

without year. Medley Wood 158 (NH); Hammarsdale, (-DA), 4 May

1995, Ward 13088 (NH, NU), Assagay, (-DC), 2 Nov. 2003, Wragg

375 (NU); Sept. 1921, Rogers 24423 (NU); Westville, (-DD), 22 Sept.

1965, Moll 2340 (NU, PRE). 2931 (Stanger): Tugela Beach, (-AB), 17

Jan. \952, Johnson 385 (NBG); Groutville, (-AC), 14 Oct. 1965, Moll

2559 (NU); The Bluff, Treasure Beach, (-CC), 6 Nov. 1982, Ellery 3

(NU); Durban, (-CC), Oct. 1883, Medley Wood 122 (PRE); 25 Oct.

1889, Medley Wood 4765 (NH); Isipingo, (-CD), May 1948, Ward 363

(NH, NU); 9 Feb. 1967, Ward 6054 (NU). 3029 (Kokstad): Kokstad,

(-AD), Dec. 1881, Tyson 1816 (SAM); Mt Currie, (-AD), 25 Nov.

1930, Goossens 341 (PRE); Weza, Ingeli slopes, (-DA), 2 Jan. 1966,

Strey 6366 (NH, PRE); Weza State Forest, (-DA), 20 Nov. 1986, Jor-

daan 857 (NH); Harding, Rooi Vaal, (-DB), 7 Jan. 1957, Taylor 5359

(NBG). 3030 (Port Shepstone): Alexandra Dist., Umgay [Umgai],

(-AD), 7 Nov. 1908, Rudatis 461 (NBG, PRE); Dumisa, Fairfield, (-

AD), 21 Oct. 1997, Arkell 324 (NH); 22 Oct. 1997, Ngwenya 1573

(NH); 10 km to Highflats from Umtentweni, (-AD), 6 Jan. 1981, Schr-

ire 573 (NH); 10 km from Highflats on road to Umzinto, (-AD), 7 Jan.

1981, Germishuizen 1812 (PRE); Dududu, The Cedars, (-BA), 5 Nov.

1992, Williams 956 (NH, PRE); Amanzimtoti, (-BB), 27 Sept. 1959,

Wilson 29 (NU); Scottburgh, (-BC), May 1954, Garbutt 15 (NU);

Pennington, (-BC), Apr. 1950, Gower 17 (NU); 20 Dec. 1960, Mauve

4098 (PRE); Ifafa, (-BC), 12 Jan. 1943, Hardley 73 (NU); Park Rynie,

(-BC), 8 Apr. 1967, Baijnath 381 (NU, PRE); Vemon-Crookes Nature

Reserve, (-BC), 11 Nov. 2001, Styles 116 (NU); Umtamvuna Nature

Reserve, (-CC), 27 Feb. 1983, Abbott 874. 875 (NH); Oribi Gorge, (-

CD), 11 Jan. 1971, Glen 328 (NU); Margate, (-CD), Feb. 1931, Rump

s.n. (NH). 3130 (Port Edward): Port Edward, Farm Blencathra, (-AA),

26 Nov. 2003, Singh & Baijnath 847 (NH).

EASTERN CAPE.— 3129 (Port St. Johns): Lusikisiki, (-BC), 24

Nov. 1964, Bayliss 2519 (NBG). 3130 (Port Edward): Mzamba, (-AA/

AB), 19 Oct.'l993, Arkell & Abbott 151 (NH). 3327 (Peddie): Fish

River Mouth, (-AC), 22 Oct. 1964, Bayliss 2458 (NBG).

7. Silene crassifolia L., Species plantarum, ed. 2,

1: 597 (1762); Sond.: 129 (1860). Type: South Africa,

[Western Cape], Strandfontein, 21 Dec. 1941, Compton

12781 [NBG, neo.!, designated by Cupido in Cafferty &

Jarvis: 1053 (2004)].

Geophytic perennial 7-30 cm tall, forming loose or

more compact mats; tuber cylindrical or ovoid; stems

prostrate or straggling, decumbent, basal woody portion

2-5 mm diam. but ( 1.0-)1. 5-2.0 mm diam. at base of

inflorescence, ± densely puberulous-scabridulous with

short, deflexed or spreading, acute hairs, rarely sub-

glabrous. Leaves cauline, lower mostly dry or withered,

oblanceolate to obovate or suborbicular, (10-) 15-50 x

5-18 mm, obtuse, apiculate or weakly uncinate, base

narrowed in lower leaves, leathery to sub-succulent, ±

densely adpressed-puberulous to thickly felted, rarely

glabrous on adaxial or both surfaces, hairs acute, mar-

gins ciliate towards base with longer, straggling hairs to

1 mm long, without evident side veins. Inflorescence a

lax or dense, (2)3-8(-10)-flowered monochasium ter-

minating main stems and branches, not evidently sepa-

rated from leafy stem, rachis 1.0-1. 5 mm diam.; bracts

smaller than leaves, unequal to subequal, ovate or obo-

vate to oblong or linear-lanceolate, puberulous to thickly

felted with appressed or erect, acute hairs, margins

densely ciliolate; pedicels 3-10(-15) mm long, rarely

up to 20 mm long in fruit. Calyx clavate or loosely

funnel-shaped in flower, ( 10-)12-18(-20) x 3^5 mm

at anthesis, strongly plicate, equally 10-veined, densely

adpressed-puberulous/felted, lobes ovate or triangu-

lar, ± 2-3 mm long, densely ciliolate. Flowers patent

at anthesis, nocturnal. Petals white to pale yellowish or

pale pink, claw ± 7(-10) mm long, rarely exserted up to

4 mm beyond calyx, narrowed below, without evident

auricles, pubescent along midrib and veins on outer sur-

face, limb bifid, 4-6 x 4-6 mm, coronal scales 0. 8-1.0

mm long. Stamen filaments slightly unequal, 6-7 mm

long, shorter series shortly included, longer series reach-

ing top of claw or exserted up to 1.5 mm. Ovaty ellip-

soid, ± 4 mm long; styles 4-5 mm long, exserted up to 2

mm. Capsule broadly ovoid or subglobose, (8-) 10- 12 x

7-10 mm, subequal to slightly longer than carpophore,

smooth; carpophore 4-8 mm long, pubescent. Seeds

1.0-1. 5 mm, reniform with deep hilar notch, flanks

flat or somewhat concave, back deeply and narrowly

grooved between two undulate wings, reddish brown,

testa striate. Chromosome number (subsp. primuliflora):

2n = 24 (Masson 1989). Flowering time: mainly Sept.-

Mar. but ± throughout the year along the east coast. Fig-

ures 3D-G, 15.

Distribution and ecology: distributed along the south-

western, southern, and southeastern coast of South

Africa, from Saldanha Bay in the Western Cape to Park

Rynie in southern KwaZulu-Natal (Figure 16). The spe-

cies is strictly coastal, growing in deep sandy soils, often

just above the high level waterline but up to 20 m a.s.l.,

typically on stablized or partially stabilized foredunes

at the edge of coastal scrub or grassland, rarely on cliffs

but evidently never on limestone or other calcareous

outcrops.

Diagnosis and relationships: Silene crassifolia is

usually readily identified by its prostrate or decumbent,

relatively robust, perennial and often branching stems,

mostly 1. 5-2.0 mm diam. at the base of the inflores-

cence, that are leafy almost to the base of the inflores-

cence, oblanceolate to obovate or sub-orbicular leaves,

somewhat inflated and strongly pleated calyx 3-5 mm

diam., and broadly um-shaped to subglobose capsules,

(8-) 10- 12 X 7-10 mm. The strictly coastal distribution,

on sandy foredunes, is also diagnostic.

Collections of subsp. primuliflora from the eastern

coast, especially the Wild Coast and KwaZulu-Natal,

may be confused with broad-leaved Silene burchellii

subsp. midtiflora and have sometimes been identified as

a coastal form of this species. S. burchellii, which gener-

ally occurs on finer-grained soils inland of the coast, is

a more slender plant, with essentially annual, mostly ±

erect stems 0.5-1. 5 mm diam. at the base of the inflores-

cence, with the foliage leaves concentrated towards the

base of the stem and well separated from the inflores-

cence by an upper portion of leafless stem, and smaller

capsules, 8-10 x 5-7 mm. This is the only region where

both species occur, and the possibility of introgression in

the coastal grasslands here cannot be discounted.

The two subspecies of Silene crassifolia recognized

here have traditionally been regarded as distinct spe-

cies but the greatly increased collections now avail-

able have led us to reconsider this interpretation. When

Sonder ( 1 860) last monographed the genus, he knew S.

crassifolia only from the Cape Flats (Blouberg and Riet

Valley [Gordon’s Bay]), and S. primuliflora from a few

collections east of Knysna (Tsitsikamma, Port Elizabeth,

and southern KwaZulu-Natal). The distinction between

the two taxa was quite clear from this sparse material:

174

Bothalia42,2 (2012)

S. crassifolia was readily diagnosed by its suborbicu-

lar or obovate, hairy leaves, ovate bracts, hirsute calyx

and relatively shorter carpophore half as long as the

capsule, and S. primuliflora by its narrower, spathulate,

scabrid leaves, ovate-lanceolate bracts, pubescent calyx,

and relatively longer carpophore ± as long as the calyx.

Later workers (Bocquet & Kiefer 1878; Masson 1989)

have maintained these distinctions but the additional col-

lections, especially those from the intervening southern

Cape coast, show that these differences are not consist-

ently maintained and that the two extremes grade into

one another, especially between Knysna and Mossel

Bay, where plants with the broad leaves and short car-

pophore of S. crassifolia develop the short pubescence

characteristic of .S', primuliflora. Further west, at Betty’s

FIGURE 15. — Silene crassifolia

subsp. crassifolia, Saldanha,

Goldblatt & Porter 13568.

A, flowering stem; B, petal;

C, gynoecium; D, seed side

view; D, seed dorsal view.

Scale bar: A, 10 mm; B, C,

2.5 mm; D, E, 0.5 mm. Artist:

John Manning.

Bay, plants have the narrow leaves of S. primuliflora

but the woolly stem and calyx of S. crassifolia, while at

Blouberg Strand some collections have the broad leaves

and bracts of S. crassifolia but the short pubescence of

S. primuliflora. Other purported differences between the

taxa, in calyx length and capsule shape, are similarly

variable across the range, with no clear discontinuity

between them. This general but incomplete association

of morphology with geography is consistent with the

rank of subspecies (Stuessy 1990).

Key to subspecies

1 a Plants mostly ± thickly felted on stems and leaves, hairs on

stem usually spreading; leaves mostly obovate to sub-

orbicular; calyx 12-15 mm long; carpophore 4-5 mm

Bothalia 42,2 (2012)

175

FIGURE 16. — Distribution of Silene crassifolia: subsp. crassifolia. •;

suhs'p. primuliflora, o.

long, ± half as long as capsule; west of Breede River

mouth 7a. subsp. crassifolia

lb Plants mostly puberulous on stems and leaves, hairs on stem

± deflexed; leaves oblanceolate to obovate; calyx mostly

(10-)15-18(-20) mm long; carpophore 5-8 mm long,

usually more than half as long as capsule; east of Breede

River Mouth 7b. subsp. primuliflora

7a. subsp. crassifolia

Plants mostly ± thickly felted with hairs on stem

deflexed or spreading. Leaves obovate to sub-orbicular,

rarely oblanceolate, ± densely pubescent adaxially and

thickly felted abaxially, rarely subglabrous adaxially and

pubescent abaxially, often sub-succulent. Calyx 12-15

mm long. Carpophore 4—5 mm long, ± half as long as

capsule. Figures 3D-F, 15.

Diagnosis', distinguished by its mostly longer, shaggy

or felted pubescence, broader leaves, and often slightly

shorter calyx, 12-15 mm long

Distribution', restricted to the southwestern and

southern coasts of Western Cape, from the coast near

Saldanha to Struisbaai (Figure 16). Plants from the Cape

Peninsula exhibit the most extreme phenotypic adapta-

tions to a maritime environment, with obovate to subor-

bicular leaves that are densely felted, especially on the

lower surface.

Additional specimens

WESTERN CAPE. — 3217 (Vredenburg): Jacobsbaai, (-DD),

25 Sept. 2009, Goldblatt, Manning & Porter 13447 (MO, NBG);

Saldanha, Tabacbaai, (-DD), 25 Sept. 2009, Goldblatt & Porter 13568

(MO, NBG). 3318 (Cape Town): Langebaan Peninsula, Schrywers-

hoek, ± 20 m, (-AA), 26 Nov. 1975, Boucher 2954 (NBG); Yzerfon-

tein, (-AC), 1 Nov. 1953, Levyns 10024 (BOL); Blouberg Strand, (-

AD), Oct. 1832, Zeyher 12 (SAM); 5 June 1940, Bond 415 (NBG); 3

Oct. 2010, Goldbatt & Manning 13600 (MO, NBG); Rietvalley [Riet-

vlei], (-AD), Nov. without year, Ecklon & Zeyher 255 (SAM). 3418

(Simonstown): Fish Hoek, (-AB), Sept. 1882, Bolus 4763 (BOL);

Kalk Bay, (-AB), Jan. 1880, Bolus 4763 (BOL); Muizenberg, (-AB),

16 May 1903, Pearson 35 (NBG); 2 Dec. 1938, Adamson 2320 (BOL);

Strandfontein, (-AB), 18 Sept. 1942, Compton 13706 (NBG); 22

Oct. 1962, Taylor 4189 (NBG, PRE); Nov. without year, Lamb 1951

(SAM); Cape Peninsula, Platboom, (-AD), 6 Nov. 1941, Compton

12297 (NBG); Seekoe River Mouth, (-BA), 16 Sept. 1980, Parsons

5 (PRE); Somerset West, Swartklip, (-BA), 15 Aug. 1953, Leistner

1113 (NBG); Macassar, dunes, (-BA), 23 Nov. 1955, Van der Merwe

42 (NBG); N of Shuster River, foredunes, 15 m, (-BA), 12 Nov. 1986,

O'Callaghan 1365 (NBG); Betty’s Bay, (-BD), 21 Nov. 1952, Parker

4833 (BOL, NBG); 28 Jan. 1970, Boucher 1099 (NBG, PRE); 25 Sept.

1996, Forrester 1143 (NBG); Rooiels, (-BD), 27 Jan. 1962, Walsh s.n.

(NBG, PRE). 3419 (Caledon): Kleinmond, (-AC), without date, De

Vos 255 (NBG); Bot River Vlei, sandy dunes, 20 m, (-AC), 12 Aug.

1982, O’Callaghan 206 (NBG); Pearly Beach, (-DA), 4 Nov. 1969,

Taylor 7425 (NBG); 9 March 1980, Raitt 447 (NBG); Agulhas, beyond

lighthouse, (-DD), 16 Nov. 1979, Taylor 10157 (NBG, PRE). 3420

(Bredasdorp): Lekkerwater, Farm Hamerkop, (-BC), 28 Nov. 1978,

Taylor 9903 (NBG, PRE).

7b. subsp. primuliflora (Eckl. & Zeyh.) J.C. Manning

& Goldblatt, comb, et stat. nov. Silene primuliflora

Eckl. & Zeyh. [as flrimulaeflora']: 32 (1834); Sond.:

129 (1860). Type: South Africa, [Eastern Cape], ‘campo

marino arenoso sinus Algoabay prope Port Elisabeth

osque fluvii Zwartkopsrivier’, Oct. [without year], Eck-

lon & Zeyher 618 (SAM, lecto.!, designated by Bocquet

& Keifer; 8 (1978); SAM, isolecto.!).

Silene primuliflora var. ciliata Fenzl. ex Sond.: 129

(1860), syn. nov. Type: not designated.

Silene vlokii D. Masson: 485 (1989). Type: South

Afriea, [Western Cape], 22 Oct. 1987, D. Masson 1225

(G, holo.!, iso.!).

Silene colorata var. ciliata Fenzl. in Drege: 222

(1844), nom. nud.

Plants mostly puberulous with hairs on stem ±

deflexed. Leaves oblanceolate to obovate, ± puberulous,

more densely so abaxially, rarely glabrous on adaxial or

both surfaces except along margins. Calyx (10-)15-18(-

20) mm long. Carpophore 5-8 mm long, mostly more

than half as long as capsule. Figure 3G.

Diagnosis', distinguished by its shorter pubescence,

generally narrower leaves and often slightly longer

calyx, (10-)15-18(-20) mm long.

Distribution', distributed along the southern and

eastern coast of South Africa, from Vermaaklikheid

just east of the Breede River Mouth along the southern

and southeastern coast into KwaZulu-Natal where it is

mainly recorded south of Durban, with one collection

from Shaka’s Rock on the North Coast northeast of Dur-

ban (Figure 16). Populations between East London and

Mazeppa Bay tend to have ± glabrous leaf blades and

were distinguished as var. ciliata by Sonder (1860) but

this is not consistent, e.g. Maguire 3675 from Kei River

Bridge and Wisura 2655 from Mazeppa Bay, whieh

include individuals that are either glabrous or variously

puberulous on one or both leaf surfaces. Plants from the

cliffs at Herold’s Bay near George described as S. vlokii

(Masson 1989) represent a dwarf form differing only in

their more compact habit, typical of plants from exposed

situations such as this, and were synonymized under S.

primuliflora by Goldblatt & Manning (2000).

Additional specimens

KWAZULU-NATAL.— 2931 (Stanger): Chaka’s [Shaka’s] Rock,

(-CA), without date, Hillary 156 (NU). 3030 (Port Shepstone): Ifafa,

(-BC), 12 Jul. 1974, Huntley 206 (NU); Park Rynie, (-BC), May 1906,

Thode 5513 (NBG); Uvongo Beach, (-CB), 4 Aug. 1967, Strey 7602

(NU, PRE); Ramsgate, (-CB), 7 July 1989, Vos & Gormley s.n. (NU);

16 Sept. 2003, Styles 1590 (NU). 3130 (Port Edward): Port Edward, (-

AA), 16 Nov. 1963, Lennox s.n (NU).

176

Bothalia 42,2 (2012)

WESTERN CAPE. — 3421 (Riversdale): Farm Koensrus near Ver-

maaklikheid. (-AC), 3 Mar, 2000, Goldblatt & Ndnni 11292A (MO,

NBG). 3422 (Mossel Bay): Great Brak River Mouth, (-AA), 13 Nov.

1981, Parsons 375 (NBG); Herold’s Bay, (~AB), 7 Jan. 1995, Victor

864 (PRE); Kleinkrans just outside Wilderness, (-BA), 19 Jan. 1943,

Compton 14333 (NBG); 6 Feb. 1944, Compton 155 73 (NBG); 7 Nov.

1979, Hugo 1915 (NBG, PRE); Wilderness, beach near river mouth,

(-BA), 23 Dec. 1987, Vlok 1900 (PRE); Wilderness, dunes E of river

bridge, (-BA), 19 Sept, 2010, Goldblatt & Porter 13548 (NBG);

Sedgefield, dunes at mouth of vlei, (-BB), 1 Dec. 1959, Middlemost

2046 (NBG). 3423 (Knysna); Knysna Heads, (-AA), 7 Nov. 1928, Gil-

lett 2168 (NBG); Fourcade 4144 (NBG); Buffalo Bay, (-AA), 4 dec.

1962, Taylor 4442 (PRE); sand dunes E of Robberg, (-AB), 1 Dec.

1943, Fourcade 6311 (NBG); Keurbooms River, (-AB), 9 Feb. 1936,

Gillen 141 7 (BOL); 28 Feb. 1 948, Price s.n. BOL24957 (BOL); 8 Mar.

1983, O’Callaghan 847 (NBG).

EASTERN CAPE. — 3129 (Port St Johns); Mkambati Nature

Reserve, between Daza and Msikaba Rivers, (-BD), 11 Dec. 1986,

Jordaan 986 (NH); Msikaba, (-BD), 5 Aug. 1972, Coleman 598 (NH);

Umgazi River Mouth, (-CB), 9 Dec. 1975, Taydor 8991 (PRE); Port

St Johns, (-DA), Dec. 1943, Brueckner & Allsopp 200 (NU); 1 Oct.

1962, Strey 4325 (NH, PRE); Coffee Bay, (-CC), Mar. 1947, Lewis

s.n. SAM63435 (SAM); Dec. 1960, Van der Shijff 5448 (PRE). 3228

(Butterworth); Bashee River Mouth, The Haven, (-BB), 17 Oct. 1966,

Gordon-Gray 58977 (NH, NU); Mendwana River Mouth, (-BC), 26

July 1965, Wood 81 (NU); Mazeppa Bay, (-BC), 15 Jun. 1973, Wisura

2655 (NBG); Kei River, (-CB), 19 Oct. 1951, Taylor 3675 (NBG);

Kei Mouth, (-CB), Jul. 1889, Flanagan 204b (PRE); 29 Mar. 1973,

Strey 11237 (MO, NU); Morgan’s Bay, (-CB), 15 Jan. 1951, Wilman

1057 (BOL); Morgan’s Bay, Double Mouth, (-CB), 19 Jan. 1979,

Hilliard & Burn 12453 (NU); Kentani Dist., coast along edge of scrub

in sea sand, (-CB), 4 Dec. 1905 & Jul. 1906, Pegler 1311 (BOL); N

side of Nxaco [River] Mouth, landward side of dune, (-CB), 12 July

1966, Ward 5732 (NH, NU, PRE). 3323 (Willowmore); Nature’s Val-

ley, (-DC), 22 Jan. 1978, Taylor 476/2 (NBG). 3325 (Port Elizabeth);

Port Elizabeth, Swartkops River Mouth, (-DC), 23 Jan. 1973 [fruit-

ing], Dahlstrand 3004 (PRE); Markhan Industrial Area, (-DC), 29

May 1973, Dahlstrand 3050 (MO, NBG, PRE). 3326 (Grahamstown);

Boknes, (-DA), Jan. 1949, Leighton 3097 (BOL, PRE); 20 oct. 1979,

Botha 2630 (PRE); Kenton-on-Sea, (-DA), Dec. 1949, Leighton 3096

(BOL, PRE); 27 Jun. 1955, Acocks 18325 (PRE); Kasouga [Kasuka]

River Mouth. (-DA), 21 Sept. 1920, Britten 2304 (PRE); 22 Sept.

1920, Britten 2343 (PRE); Kariega River Mouth, (-DA), 6 Dec. 1977,

Hilliard & Burn 10872 (E, K, MO, NU); Kowie, (-DB), 8 Feb, 1921,

Britten 2689 (PRE); 8 Jul. 1931, Levyns 3778 (BOL); Port Alfred, (-

DB), 25 Nov. 1888, Galpin 343a (PRE); Jan. 1907, Potts 187 (BOL);

Jan. 1916, Tyson s.n. PRE12592 (PRE). 3327 (Peddie); Hamburg, (-

AB), 28 Dec. 1931, Gemtnell s.n. (PRE); [Great] Fish River Mouth,

(-AC), Nov. [without year], MacOwan 737 (BOL); East London, (-

BB), Feb. 1888, Thode STE6598 (NBG); Jan. 1914, Potts 1783 (BOL);

Feb. 1920, Page s.n. BOLI6978 (BOL); Apr. -Jun. 1961, Bokelmann

2 (NBG); Gonubie Beach, (-BB), Jan. 1953, Peacock s.n. SAM67466

(SAM); Gonubie Mouth, (-BB), 4 Oct. 1942, Acocks 9137 (PRE);

Shelly Beach, (-BB), Dec. 1952, Peacock s.n. SAM67467 (SAM); 27

Jan. 1979, Hilliard & Burn 12408 (NU). 3424 (Humansdorp); Groot

River Mouth, (-AA), Mar. 1910, Fourcade 6676 (BOL); 29 Dec. 1926,

Phillips 1204 (BOL); Jan. 1927, Duthie 547 (NBG, SAM); 8 Apr.

1981, Parsons 195 (NBG); Jeffrey’s Bay, (-BA), 17 Dec. 1956, Tay-

lor 5147 (NBG). 3425 (Skoenmakerskop); Skoenmakerskop, (-BA),

7 Sept. 2002, Stevn 92 (PRE); Cape Recife, (-BA), 1987, D. Masson

1230 (VKE).

8. Silene mundiana Eckl. & Zeyh., Enumeratio

plantarum africae australis extratropicae: 32 (1834);

Sond.: 127 (1860) [as ‘’mundtiana'}. Type: South Africa,

[Eastern Cape], ‘iter frutices ad Paardekop prope sinum

Plettenbergsbay [Plettenberg Bay]’, without date, Mund

s.n. (SAM, lecto.!, designated by Boequet & Kiefer: 8

( 1 978); S, isolecto.).

Geophytic perennial 7-30 cm tall, forming loose,

tangled mounds or more compact mats; stems sprawl-

ing or straggling, decumbent, highly and often closely

branched, slender, basal woody portion 1-2 mm diam.

but 0. 5-1.0 mm diam. at base of inflorescence, closely

puberulous-scabridulous with very short, deflexed, acute

hairs. Leaves cauline, lower mostly dry or withered,

oblanceolate to linear-oblanceolate, 7-15 x 1-3 mm,

apiculate or weakly uncinate, base narrowed in lower

leaves, puberulous along lower midline and margins,

and sometimes also basally, hairs acute, margins ciliate

towards base with longer, straggling hairs to 1 mm long,

without evident side veins. Inflorescence a (l)2-flowered

monochasium terminating main stems and branches,

rachis filiform, 0.2-0. 5 mm diam., not extending much

beyond vegetative growth; bracts smaller than leaves,

subequal, linear, thinly puberulous with appressed,

acute hairs, margins densely ciliolate; pedicels filiform,

3- 10(-15) mm long. Calyx clavate in flower, 9-11 x

2. 5-3.0 mm at anthesis, equally 10-veined, densely

adpressed-puberulous/scabridulous, lobes ovate or tri-

angular, ± 1.5 mm long, densely ciliolate. Flowers pat-

ent at anthesis, ?noctumal. Petals white (?or pale pink),

claw ± 6 mm long, narrowed below, without evident

auricles, pubescent along midrib and veins on outer

surface, limb bifid, 4-5 x 3^ mm, coronal scales 0.5-

0.8 mm long. Stamen filaments slightly unequal, 6-7

mm long, shorter series shortly included, longer series

shortly exserted. Ovary ellipsoid, ± 3 mm long; styles

4- 5 mm long, exserted up to 1 mm. Capsule broadly

ovoid, 6x4 mm, slightly longer than carpophore,

smooth; carpophore ± 4 mm long, pubescent. Seeds 0.8-

1.0 mm, reniform with deep hilar notch, flanks flat or

somewhat concave, back deeply and narrowly grooved

between two undulate wings, reddish brown, testa stri-

ate. Flowering time: Sept.-Oct.(-Nov.). Figure 3H, L

Ecology and distribution: originally described from

Plettenberg Bay, where it is still known only from the

type and thus possibly extinct locally, the species is now

also known to occur near Bredasdorp, more than 200

km to the west, in the De Hoop Nature Reserve imme-

diately west of Potberg (Figure 10). Here it is restricted

to limestone outcrops near the coast, either fringing sea-

sonal vleis or directly facing the sea. Plants growing on

exposed cliffs display the typical maritime adaptations

of a more compact habit, forming ± prostrate closely

leafy mats, and with smaller, thicker leaves than those

sheltered in potholes, which are more diffuse, forming

loose cushions in the crevices. The species appears to be

locally common where it occurs at De Hoop.

Diagnosis and relationships: one of the least known

of the South African species, represented in herbaria by

just a handful of specimens.

The highly reduced inflorescences of Silene mun-

diana, not immediately identifiable as monochasia,

led Sonder (1860) to place the species provisionally

[‘doubtfully’] in sect. Elisanthe but its geophytic habit,

eglandular vestiture, ovate calyx lobes, and especially

the winged seeds place it correctly in sect. Fruticu-

losae. Here it is distinguished by its densely and closely

branched habit, small leaves 7-15 x [-3 mm, and ± fili-

form branches with just 1 or 2 flowers per inflorescence.

Plants form small mats or low tangled mounds, with the

numerous inflorescences extending only shortly above

the vegetative growth.

Conservation note: the restricted occurrence of the

species and its possible extinction around Plettenberg

Bay render it vulnerable.

Bothalia 42,2 (2012)

177

Additional specimens

WESTERN CAPE. — 3420 (Bredasdorp): De Hoop farm in mud

of dried up vlei, (-AC), 1971 [without month]. Van der Merwe 1871

(NBG); De Hoop, Potberg Nature reserve, limestone cliffs along vlei

2 km NW of De Hoop residence, (-AD), 16 Oct. 1978, Burgers 1358

(NBG); De Hoop Nature Reserve near Rest Camp, in exposed rock

crevices facing coastal winds, (-AD), 22 Oct. 1980, Mauve. Reid &

Wikner 30 (NBG).

Silene sect. Dipterospermae (Rohrb.) Chowdlniri,

Notes from the Royal Botanic Garden, Edinburgh 22:

248 (1957). Silene [unranked] Dipterospermae Rohrb.:

69 (1869). Lectotype: Silene colorata Poir., designated

by Chowdhuri: 248 (1957).

Annuals, eglandular pubescent. Leaves linear to obo-

vate, lacking apparent lateral veins; bracts herbaceous.

Flowers in monochasia with axis simple or forked

below, ± erect, diurnal, vespertine or cleistogamous;

anthophore well-developed, pubescent. Calyx pubescent,

clavate, 10-veined, without conspicuous anastomos-

ing venation, lobes ovate, obtuse. Petals: claw scabrid

along midvein above, not auriculate, limb bifid with

linear lobes; coronal scales present. Ovaty partially sep-

tate. Seeds reniform with deep hilar notch, flanks flat or

somewhat concave and smooth or striate, back deeply

and narrowly grooved between two undulate peripheral

wings.

± 9 spp., circum-Mediterranean to Arabia and W

Pakistan, South Africa.

Although Chowdhuri (1957) placed Silene aethio-

pica [as S. clandestina] in sect. Scorpioideae, it is highly

anomalous there in its peripherally winged seeds (Fig-

ure 3J-M). These, together with the annual habit, locate

it firmly in sect. Dipterospermae. Seeds of this type are

otherwise characteristic of sect. Fruticulosae (Figure

3A-I) and both groups also share a clavate calyx with

ovate calyx lobes. The two sections are essentially sepa-

rated on the basis of an annual vs. a perennial habit, and

their exact relationship requires further study. It is espe-

cially suggestive that Gilbert (2000) notes that forms of

the usually perennial S. burchellii (sect. Fruticulosae)

from the drier parts of Ethiopia are often ephemerals

with an evident similarity to the annual 5. colorata (sect.

Dipterospermae).

9. Silene aethiopica Burm., Flora indica: cui accedit

series zoophytorum indicorum, nec non Prodromus flo-

rae capensis: 13 (1768). Type: South Africa, without

precise locality or collector, Burman s.n. Herb. Burm.

G00301755 (G, lecto. — digital image!, designated

here). [Of the two available syntypes, G00301754 and

G00301755, the latter is selected as lectotype as it is a

closer match to the protologue in its much branched

stem and oblanceolate leaves].

Annual, 7-30(-35) cm tall; stems erect or decum-

bent, mostly branching from near base, 0.75 mm diam.

at base but 0.5-1. 5 mm diam. at inflorescence, closely

puberulous with very short, deflexed hairs, rarely gla-

brescent. Leaves linear or narrowly oblanceolate to

spathulate-oblanceolate, becoming narrower distally,

15-60 X 2-5(-10) mm, apiculate or obtuse, base nar-

rowed in lower leaves, ± pubescent with spreading or

± appressed hairs, rarely glabrescent or glabrous apart

from margins, margins towards base ciliate with hairs to

1 mm long, without evident side veins. Inflorescence a

secund, spike-like monochasium terminating main stems

and branches, 2-8-flowered; bracts smaller than leaves,

unequal, fertile bract linear to awl-shaped, densely

appressed-puberulous with acute, eglandular hairs; pedi-

cels up to 25 mm long in lower flowers and elongat-

ing to 35 mm long in fruit, (2-)6-10 mm long in upper

flowers, erect in fruit. Calyx clavate in flower, 10-15(-

18) X 2-5 mm at anthesis, equally 10-veined, densely

adpressed-puberulous, lobes ovate to lanceolate, ± 2 mm

long, densely cilio-late. Flowers suberect at anthesis,

unscented or sweetly scented of coconut-vanilla, noc-

turnal. Petals white to pink with maroon reverse, claw

6-8 mm long, narrowly oblanceolate, without auricles,

limb bifid or emarginate, (l-)3-8 mm x 2-6 mm, coro-

nal scales 0. 5-1.0 mm long. Stamen filaments slightly

unequal, 4. 5-7.0 mm long, shorter series included up to

2 mm, longer series ± reaching top of claw. Ovary ellip-

soid, ± 3 mm long; styles 3-5 mm long, ± reaching top

of claw. Capsule ovoid, (6-)8-9 x 3^ mm, ± as long

as to 2x as long as carpophore; carpophore (3-)4-9

mm long, pubescent. Seeds 0.8-1. 5 mm, reniform with

deep hilar notch, flanks flat or somewhat concave, back

deeply and narrowly grooved between two undulate

wings, reddish brown, testa striate. Chromosome number

(subsp. maritima): n = 12 (Bocquet 1977). Flowering

time: (Jul.)Aug.-Oct.(-Nov.). Figures 3J-M, 17.

Distribution and ecology’: widely distributed through

the southern African winter rainfall region, mainly on

the coastal lowlands, from the southern Richtersveld

southwards to Stilbaai and inland to the Tankwa Karoo

and Faingsburg, with a disjunction to Port Elizabeth

in the east but evidently absent from the Fittle Karoo

(Figure 18). The species is mostly encountered in well-

drained, coarse-grained soils of a wide variety of types,

ranging from acidic sands and gravels derived from

sandstone and granite to basic limestones and calcare-

ous sands, very rarely in fine-grained clays, occurring on

sandy flats, in dry river courses, around salt pans and on

limestone, or rarely dolerite, ridges.

Diagnosis and relationships: the only annual species

of Silene native to southern Africa, S. aethiopica is dis-

tinguished from the introduced annual species, S. gallica

and S. nocturna, by its wholly eglandular pubescence —

the stems with short, deflexed hairs and the clavate calyx

covered with short, appressed hairs — and by its discoid,

peripherally winged seeds. Above ground it is superfi-

cially similar to the perennial 5. burchellii (sect. Fruticu-

losae), with similar vestiture, calyx, and seeds and the

two species can be confused in the southwestern parts

of the Western Cape where their ranges overlap. Well-

grown specimens of S. aethiopica with branched stems

are unmistakable but unbranched specimens without

roots may pose problems. Calyx length is a useful dis-

criminating feature in the region of overlap, with the

calyx of S. aethiopica typically 10-15 mm long, except

in subsp. longiflora from the Cold Bokkeveld, where it

is 15-18 mm long. In contrast, the more common forms

of S. burchellii in the region, subsp. pilosellifolia, have a

calyx 18-25 mm long, although clayx length is shorter,

15-20 mm long, in subsp. burchellii in the extreme

southwestern coastal areas. Ecologically, S. aethiopica

178

Bothalia 42,2 (2012)

FIGURE 17. — Silene aethiopica

subsp. aethiopica: A, typical

form from Tulbagh, no vouch-

er; B, typical form from Gor-

don’s Bay, no voucher; C-K,

maritime form from Blouberg

Strand, Goldblatt & Manning

1 3601. A, D, flowering plants;

B, flowering stem; C, stem

segment; E, calyx laid out;

F, petal; G, androecium laid

out; H, gynoecium; I, capsule

before dehischence with calyx

removed; J, seed side view; K,

seed dorsal view. Scale bar: A,

B, D, I, 10 mm; C, 1 mm; E, 5

mm; F-H, 2.5 mm; J, K, 0.5.

Artist: John Manning.

prefers well-drained, sandy soils, whereas S. burchellii

is mostly found in stony places or in finer-grained clays.

The two taxa are, however, best separated by habit. S.

aethiopica is an annual species with a slender tap-

root that is invariably present in herbarium specimens,

which are always pulled from the ground with the main

root intact. S. burchellii, in contrast, is a perennial with

annual flowering stems but a persistent, woody stem

base, developing a persistent, short or long rootstock

that is swollen and tuber-like distally, although the swol-

len terminal portion is very rarely present in herbarium

specimens.

As in other species of Silene, S. aethiopica is variable

in stature, leaf morphology and flower size, notably in

the length of petal limbs, carpophore and calyx. Much

of the variation is either ecological or recurrent through

the range of the species but populations from the south-

ern Knersvlakte and the Olifants River valley are con-

sistently less pubescent than usual, with the leaf blades

either glabrous, apart from the margins, or glabrescent,

and the stems often glabrescent as well. Large- and

small-flowered forms occur throughout the range of the

species, sometimes within single populations.

Strand forms of Silene aethiopica from the south-

western coast (Figure 17D), distinguished from more

inland plants (Figure 17A, B) by the typical maritime

adaptations of a more spreading habit, broader and more

fleshy leaves, and fleshy caljoc with distinctly bulg-

ing nerves, were described as a separate species under

the name S. dewinteri (Bocquet 1977). The distinc-

tion between the two forms is not absolute, however,

and some collections comprise material that could be

assigned to both taxa (eg. Goldblatt & Porter 13509).

We visited the type locality of S. dewinteri at Table

View and confirm Bocquet ’s observations regarding the

texture of the leaves and calyx but not of the habit —

all of the plants that we encountered were erect, possi-

bly because our visit to the site was earlier in the sea-

son than Bocquet’s. More significantly, however, our

transect across the dunes passed through a gradation

from typical S. aethiopica on the landward side to S.

dewinteri on the seaward side. We encountered similar

Bothalia 42,2 (2012)

179

FIGURE 18. — Distribution of Silene aethiopica: subsp. aethiopica, •;

subsp. longiflora, o.

morphological dines at Jacobsbaai and at Elands Bay.

Although plants at the two extremes of the ecological

spectrum are certainly very different in appearance, they

grade into one another in the ecotone, leading us to con-

clude that S. dewinteri represents a ± distinct ecotype

rather than a separate species. This interpretation is also

implicit in Bocquet’s (1978) subsequent description of 5.

dewinteri as a ‘sand-dune ecotype’.

That there is some genetic basis for the maritime form

seems to be borne out by Bocquet’s (1977) observation

that the greater fleshiness of the plants was retained in

cultivated plants grown from seed. It is not clear, how-

ever, whether this variant represents recurrent selection

for a maritime genotype, a situation that is readily envis-

aged in a self-fertile species. In any event, we do not

recognize this maritime form at any taxonomic rank. An

investigation of the population genetics of the species is

necessary before the taxonomic and evolutionary status

of these variants can be accurately assessed.

A more clear-cut morphological disjunction is evident

in populations from the Cold Bokkeveld and adjacent

Cedarberg-Bidouw Mtns, which have markedly longer

calyces (and carpophores) than usual. We describe this

variant as subsp. longiflora.

Nomenclatural note: The identity of Silene aethi-

opica Burm. (1768) has remained uncertain until now.

Based on unlocalised material from the ‘Cape of Good

Hope’ and one of the earliest names published in Afri-

can silenes, the published diagnosis caule ramoso, flori-

bus subspicatus, petalis bfldis obtusis, foliis lanceolatis

viscidis includes two characteristics not combined in

any southern African species, viz. ‘viscid leaves’ and a

‘subspicate’ inflorescence. Thunberg (1823) treated the

species as a synonym of what he called S. noct flora (a

European species) but his application of this name in

the southern African context actually encompasses the

newly described S. rigens, a cymose-flowered species

in sect. Elisanthe, and the basis for his interpretation

of S. aethiopica is unclear. Others dealing with Silene

ignored the name, including Otth (1824), Ecklon &

Zeyher (1834), and Sonder (1860) in his account of the

genus for Flora capensis. Williams (1896), however, in

his revision of Silene, cited S. aethiopica as an uncertain

synonym of the later S. biirchellii, and the African Plants

Database (2011 version) follows this lead. This tenta-

tive identification of Burman’s species is clearly based

on the ‘subspicate’ nature of the infloresence. It is most

fortunate, therefore, that Nieolas Fumeaux at the Bur-

man Herbarium has succeeded in locating two sheets

bearing the name S. aethiopica, which clearly constitute

original material of the species and are thus syntypes

of the name. One of the sheets (G00301754) also bears

the inscription Lychnoides aethiopica angustifolia fl.

incarnatis, petalis acutis from John Ray’s (1704) His-

toire plantariim, and the other (G00301 754) bears an

additional manuscript name and diagnosis for Lychnis

monomotapensis. Both collections represent the species

known until now as Silene clandestina, and G0030175

was in fact identified under this name by Daniel Mas-

son in 1991. The name S. aethiopica Burm. (1768) thus

takes priority over S. clandestina Jacq. (1791), which

becomes a later synonym. Burman’s description of the

leaves of this species as viscid must be assumed to be

an error as the type material does not show this feature.

Curiously, the same specimen had earlier been deter-

mined as S. dewinteri Bocquet (1977) by Daniel Boc-

quet in 1976, who should therefore have taken up the

epithet aethiopica for this taxon were it to be recog-

nized at species level, which we do not. In any event, we

regard the Burman specimen to represent a slightly luxu-

riant individual (possibly cultivated) of typical S. aethi-

opica and not the maritime "dewinteri' form.

Although the source of Burman’s specimens is not

given, other collections of South African material in the

Burman herbarium are known to have been made by

[Heinrich] Oldenland, who was was employed by the

Dutch East India Company at the Company Gardens in

Cape Town in the late seventeenth century. Ray (1704)

certainly used material collected by Oldenland in the

preparation of his Historia plantariim, and his mention

of the species is a clear indication that it had reached

Europe by the early eighteenth century.

Key to subspecies

la Caljrx 10-15 mm long; carpophore 3-8 mm long, shorter

than to as long as capsule 9a. subsp. aethiopica

lb Calyx 15-18 mm long; carpophore ± 10 mm long, slightly

longer than capsule; Cold Bokkeveld 9b. subsp. longiflora

9a. subsp. aethiopica

Silene clandestina Jacq.: Ill (1791 [as ‘1789’]), syn.

now; Sond.: 127 (1860). Type: South Africa, without

precise locality, illustration in Jacq., Collectanea pi. 3,

fig.3 (1791 [as ‘1789’] (lecto., designated here).

Silene cerniia Thunb.: 81 (1794), syn. now Silene

biirchellii var. cerniia (Thunb.) Williams: 76 (1896).

Type: South Africa, [Western Cape], without precise

locality or date, Thunberg s.n.. Herb. Thunb. 10714

(UPS-THUNB — microfiche!, lecto., designated here).

[This specimen is selected as the most complete of the

three under this name in the Thunberg Herbarium ( UPS-

THUNB10713, 10714 & 10715), clearly showing the

annual habit of the species]

180

Bothalia42,2 (2012)

Silene linifolia Willd.: 473 (1809) {fide Sond. (I860)],

syn. nov. Type: not designated and identity uncertain.

Silene recta Bartl.: 623 (1832) [fide Sond. (I860)],

syn. nov. Type: South Africa, [Western Cape], ‘auf dem

Tafelberge auf dem zweiten Hohe’, without date, Ecklon

& Zeyher s.n. (not located).

Silene crassifolia var. angustifolia Bartl.: 623 (1832),

syn. nov. Type: South Africa, [Western Cape], ‘beim

obersten Blockhause am Teufelsberge’, without date,

Ecklon & Zeyher s.n. (not located).

Silene constantia Eckl. & Zeyh.: 32 (1834), syn. nov.

Type: South Africa, [Western Cape], ‘collium capen-

sium prope Constantiam et Hottentotsholland’, Ecklon &

Zeyher 251 (SAM, lecto.!, designated here). [Bocquet’s

(1977) contention that Ecklon & Zeyher (1834) automat-

ically rendered this name illegtimate and superfluous by

citing S. crassifolia var. angustifolia Bartl. in synonomy

is incorrect as names have priority only at their rank: S.

constantia is therefore a legitimate name at species rank,

with its own type (McNeill et al. 2006: Art. 1 1 .2)].

Silene clandestina var. minor Sond.: 128 (1860), syn.

nov. Type: not designated.

Silene clandestina var. major Sond.: 128 (1860), syn.

nov. Type: not designated.

Silene dewinteri Bocquet, Bothalia 12: 309 (1977),

syn. nov. Type: South Africa, [Western Cape], Table

View, 9 Nov. 1975, Bocquet 17774 (ZT, holo.!).

Plants ± lax or compact. Stems erect or decumbent.

Leaves linear to oblanceolate, 2-5 mm wide. Calyx

10-15 mm long. Carpophore (3-)4-8 mm long, half as

long to as long as capsule. Figure 17.

Diagnosis: distinguished from subsp. longiflora by

the shorter calyx, 10-15 mm long.

Distribution: throughout the range of the species but

replaced on the Cold Bokkeveld and Cedarberg-Bidouw

Mtns by subsp. longiflora (Figure 18). The maritime

ecotype, characterized by a compact habit, sub-succu-

lent leaves 3-10 mm wide, ± fleshy calyx, and consist-

ently short carpophore, 4-5 mm long, is restricted to the

seaward side of coastal dunes above the high-tide mark

in the extreme southwestern Western Cape between

Saldanha Bay and Onrus River Mouth, in stabilized or

semi-stabilized, calcareous sands. The absence of this

variant along the coast to the north and east may be due

to the lack of suitable sandy beaches there.

Additional specimens

NORTHERN CAPE. — 2917 (Springbok): between Kawarass and

Lekkersing, 250 m, (-AA), 4 Sept. 1925, Marlolh 12437 (NBG); turn-

off to Komaggas on Koingnaas-Kleinzee Road, (-CC), 1 1 Aug. 2007,

Be.'iter 7895 (PRE); Farm Zonnekwa, 200 m, (-CD), 7 Oct. 1986, Le

Roux & Lloyd 519 (NBG); Spektakel, (-DA), Sept. 1883, Boltis 6667

(BOL); Hester Malan Reserve, (-DB), 3 Oct. 1974, Riisch & Le Roux

836 (PRE); 26 Aug. 1985, Struck 145 (NBG); Farm De Draay, (-DB),

29 Aug. 1990, Le Roux 4122b (NBG); Groot Vlei, (-DC), 7 Sept.

1945, Compton 17296 (NBG). 2918 (Gamoep): between Gamoep and

Aggenys close to Vaalheuwel Farm, (-DC), 19 Sept. 2002, Steyn 271

(PRE). 3017 (Hondcklipbaai): Farm Koingaas, 50 m, (-AB), 27 Aug.

1986, Le Roux & Lloyd 354 (NBG); Hondeklip Bay, (-AD), Oct. 1924,

Rillans 17942 (BOL); 11 miles [17.6 km] E of Hondcklipbaai on road

to Garies, (-AD), 1 1 Sept. 1970, Thompson 1089 (NBG); Kookfontein,

(-BA), Aug. 1883, Bolus 31659 (BOL); ± 2 km W of Sarisam home-

stead, (-DA), 2 Aug. 2006, Bester 7029 (PRE). 3018 (Kamiesberg):

Platbakkies, (-BC), 10 Sept. 1976, Thompson 2866 (PRE); Langkloof,

N of Doringkraal, (-CA), 23 Sept. 2010, Goldblatt 135 73A (MO,

NBG). 3119 (Calvinia): Niewoudtville Waterfall, (-AC), Sept, 1930,

Lavis s.n. BOL198I8 (BOL); Nieuwoudtville Reserve, (-AC), 7 Sept.

1983, Perry & Snijman 2279 (NBG). 3220 (Sutherland): Farm Berg-

sigt, (-DA), 7 Sept. 1988, Crosby 965 (PRE).

WESTERN CAPE. — 3118 (Vanrhynsdorp): Draaihoek Farm, 4

km SW of Vredendal, 70 m, (-CB), 27 Aug. 1986, Hilton-Taylor 1152

(NBG); Troe-troe Farm, 3 km from Vanrhysndorp on Nieuwoudtville

road, (-DB), 3 Aug. 1977, Le Roux 2007 (NBG); Widouw Farm, 15

km S of Van Rhynsdorp, 500' [150 m], (-DB), 9 Sept. 1976, Thomp-

son 2826 (NBG, PRE); 22 Sept. 1985, Zietsman & Zietsman 1157

(PRE); Windhoek Farm at foot of Gifberg, (-DC), 22 Sept. 191, Pear-

son 6760 (BOL); Doom River Mouth, (-DC), 22 Jul. 1941, Compton

11037 (BOL, NBG); Brandewyn River, (-DD), 26 Aug. 1950, Barker

6588 (NBG); Nardouw Mtns S of Bulshoek Barrage, (-DD), 2 Sept.

1977, Goldblatt & Manning 10719 (MO, NBG). 3217 (Vredenburg):

limestone hill N of Saldanha, (-DD), 30 Sept. 2009, Goldblatt, Man-

ning & Porter 13469 (MO, NBG); dunes N of Jacobsbaai, (-DD), 9

Sept. 2010, Goldblatt & Porter 13509 (MO, NBG); Saldanha Penin-

sula, Tabacbaai, dunes along coast, (-DD), 22 Sept. 2010, Goldblatt &

Porter 13569 (MO, NBG, PRE). 3218 (Clanwilliam): Wadrif Soutpan,

(-AB), 14 Sept. 1984, O’Callaghan & Van Wyk 96 (NBG, PRE); 9-10

km S of Lambert’s Bay, (-AB), 14 Sept. 2010, Goldblatt & Porter

13533, 13534 (MO, NBG); between Leipoldtville and Elands Bay, (-

AD), Oct. 1947, Zinn s.n. SAM63443 (SAM); Elands Bay, sandy coast

below bushman cave, (-AD), 14 Sept. 2010, Goldblatt & Porter 13535

(MO, NBG); hills around Clanwilliam, (-BB), Jul. 1897, Leipoldt 594

(NBG, SAM); Olifants River Dam, (-BB), 10 Sept. 1949, Barker 5736

(NBG); near Paleisheuwel, (-BD), 5 Sept. 1954, Levyns 10162 (BOL);

Kriedouwkrantz on road to Algeria Forest Station, (-BD), 7 Sept.

1976, Hugo 429 (NBG); N of Velddrift, 10 m, (-CC), 15 Oct. 1986,

O'Callaghan 1220 (NBG); Salt pans near Zoutkloff, (-CD), 10 Sept.

1949, Barker 5803 (BOL, NBG); Piketberg Dist., Het Kruis, (-DA),

22 Sept. 1940, Compton 9519 (NBG). 3219 (Wuppertal): Driefontein

Farm, 4 km SW of Doringbos, 200 m, (-AA), 2 Oct. 1986, Hilton-

Taylor 1858 (NBG); Boontjieskloof Farm E of Pakhuis Pass, ± 1 500'

[450 m], (-AA), 4 Sept. 1976, Taylor 9319 (NBG, PRE); Heuningvlei,

Groot Koupoort, ± 3150' [945 m], (-AA), 11 Oct. 1975, Kruger 1696

(NBG); Alpha Fami, (-AA), 20 Jul. 1941, Compton 10934 (NBG);

Algeria Forest Station, (-AC), 8 Sept. 1997, Van Rooyen, Steyn & de

Villiers 710 (NBG); Cedarberg Forest reserve, Langmg, 1 000' [300

m], (-AC), 12 Sept. 1982, Viviers 581 (NBG); Tankwa Karoo National

Park, W end, (-BA), 2 Aug. 2007, Bester 7682 (NBG). 3220 (Suther-

land): Syferwater Farm, 1 212 m, (-AA), 18 Sept. 2006, Rosch 558

(NBG). 3318 (Cape Town): Langebaan, (-AA), 5 Sept. 1971, Axelson

486 (NBG); around pan SE of Jakkaisfontein, 60 m, (-AD), 24 Sept.

1996, Low 2831 (NBG); Melkbosstrand, (-AD), 24 Sept. 1966, Dahl-

strand 1066 (PRE); 30 Sept. 2010, Goldblatt & Manning 13598 (MO,

NBG); Bok Point, (-CB), 14 Sept. 1940, Barker 779 (NBG); Bok Bay,

(-CB), 14 Sept. 1940, Compton 9389 (NBG); Robben Island, (-CD),

Oct. 1932, Adamson s.n. (BOL); 20 Aug. 1943, Walgate 512 (NBG);

19 Nov. 1987, Van Jaarsveld & De Lange 9546 (NBG); Blaauwberg

[Blouberg], (-CD), Sept. 1954, Stokoe s.n. SAM67560 (SAM); Blou-

berg Strand, dunes S of Dolphin Hotel, (-CD), 3 Oct. 2010, Goldblatt

& Manning 13601 (MO, NBG); Paarden Eiland, Klein Zoar Vlei,

1 m, (-CD), 8 Oct. 1979, Linder s.n. (BOL); Milnerton, (-CD), 17

Sept. 1913, Phillips s.n. NBG103292 (NBG); ‘in leter montis tabu-

laris’, (-CD), Oct. 1873, Bolus 2718 (BOL); ‘prope Rondebosch’,

(-CD), Oct. 1878, Bolus 2718a (BOL); Bergvliet Farm, (-CD), 1-2

Nov. 1915, Purcell s.n. (SAM); Van Kampsbay [Camps Bay], without

date, Zeyher s.n. (BOL); Camps Bay, (-CD), 13 Oct. 1956, Cassidy

51 (BOL, NBG); W slopes of Table Mtn, (-CD), 5 Jul. 1936, Adam-

son 970 (PRE); Malmesbury Dist., near Pella, Burgers Post Farm,

200 m, (-DA), 9 Oct. 1979, Boucher & Shepherd 4764 (NBG, PRE);

Cape Flats Nature Reserve, University of Western Cape campus, ± 200

m, (-DC), 9 Oct. 1980, Low 1095 (NBG); Cape Flats near Tygerberg

Station, (-DC), 15 Oct. 1927, Smuts 2490 (NBG). 3319 (Worcester):

Tulbagh Waterfall, (-AC), Oct. 1928, Levyns 2566 (BOL); Laken

Vlei, (-BC), 19 Oct. 1941, Compton 12076 (NBG); Worcester, Riv-

erside, (-CB), 11 Sept. 1962, Walters 878 (NBG); Robertson, (-DD),

25 Sept. 1935, Lewis s.n. BOL31663 (BOL). 3320 (Montagu): Laings-

burg Dist., [Matjiesfontein], Fisantekraal Valley, (-BC), 7 Nov. 1948,

Compton 21152 (NBG); Montagu, (-CC), 5 Jul. 1938, Levyns 6457

(NBG). 3418 (Simonstown): Llandudno, (-Ab), 26 Sept. 1928, Hutch-

inson 535 (BOL, K); Hout Bay, (-AA), 20 Sept. 1941, Compton 11749

Bothalia42,2 (2012)

181

(NBG); Red Hill, (-AB), 6 Oct. 1962, Taylor 4128 (NBG); Clov-

elly, (-AB), 22 Oct. 1940, Adamson s.n. (BOL); Noordhoek, (-AB),

1 Nov. 1944, Barker 3293 (NBG); Muizenburg, (-AB), Dec. 1894,

Guthrie s.n. (BOL); Gifkommetjie turnoff. Cape of Good Hope Nature

Reserve, (-AD), 6 Oct. 1971, Taylor 7952 (NBG); Buffels Bay, (-AD),

8 Oct. 1966, Taylor 6949 A (PRE); Cape Flats, Swartklip, ± 170' [51

m], (-BA), 6 Sept. 1972, Taylor 8171 (NBG, PRE); Penhill, Eerste-

rivier, 50 m, (-BA), 24 Oct. 1984, Raitt 5431 (NBG); Gordon’s Bay,

(-BB), Dec. 1901, Bolus s.n. (BOL); Strand/Macassar, Somchem, 5 m,

(-BB), 28 Aug. 1995, Low 2462 (NBG). 3419 (Caledon); Onrus River,

(-AC), 28 Sept. 1958, Williams 57 (NBG). Kleinmond, (-AC), 15 Oct.

1949, De Vos 1489 (NBG); Hermanus, Riviera, (-AC), 5 Oct. 1916,

Purcell s.n. (SAM); road from Stanford to Gansbaai, (-CB), 21 Sept.

1938, Gillett 4294 (BOL); Baviaansfontein Farm, ± 3.5 miles [5.6 km]

E of De Kelders, (-CB), 27 Sept. 1962, Taylor 4106 (NBG); Uilkraals

River, 2 m, (-CB), 3 Nov. 1987, O'Callaghan 3/ll/2e (NBG). 3420

(Bredasdorp): De Hoop, Potberg Nature Reserve, (-AD), 2 Aug. 1979,

Burgers 2145 (NBG). 3421 (Riversdale): Jongensfontein Farm, Still-

bay, (-AD), 23 Aug. 1978, Bohnen 3995 (NBG); Melkhoutfontein, (-

BD), 7 Oct. 1897, Galpin 3775 (PRE).

EASTERN CAPE. — 3325 (Port Elizabeth): Zwartkopsrivier, (-

DC), Sept, without year, Zeyher 1961 Sll-24180 (S); Swartkops River

near Port Elizabeth, (-DC), 28 Aug. 1947, Rodin 1064 (BOL).

9b. subsp. longiflora J.C. Manning & Goldblatt,

subsp. nov.

TYPE. — Western Cape, 3219 (Wuppertal): Swartrug-

gens, Knolfontein Farm, 60 km NE of Ceres, 1 184 m,

(-DC), 25 Oct. 2006, 1. Jardine & C. Jardine 562 (NBG,

holo.).

Plants ± lax. Stems erect. Leaves narrowly oblanceo-

late, 3-5 mm wide. Calyx 16-18 mm long. Carpophore

± 10 mm long, slightly longer than capsule.

Diagnosis: distinguished by its longer calyx, 16-18

mm long.

Distribution and ecology’: restricted to the Cold

Bokkeveld and Cedarberg-Bidouw Mtns, occurring in

sandy soils on sandstone slopes at mid-altitudes (Figure

18).

Additional specimens

WESTERN CAPE. — 3219 (Wuppertal): pass into Biedouw Valley,

6.2 miles [10 km] from Clanwilliam-Calvinia road, 1 500' [450 m],

(-AA), 27 Aug. 1967, Thompson 351 pp. (NBG); Driehoek Valley, (-

CA), 3 Oct. 1952, Esterhuysen 20573 (BOL); Swartruggens, Knolfon-

tein Farm, 60 km NE of Ceres, 1 185 m, (-DC), 7 Oct. 2005, Jardine

& Jardine 230 (NBG).

Silene sect. Silene

Annuals, with eglandular and mostly glandular

hairs. Leaves: lowermost obovate-spathulate. Flowers

in monochasia, with axis simple or 1 -3-forked below,

spreading or erect, vespertine, diurnal or cleistogamous;

anthophore short, pubescent. Calyx pubescent to hir-

sute, with or without glandular hairs, 10-veined, with

or without anastomising venation, tubular to fusiform in

flower, tapering apically in fruit. Petals: claw glabrous,

not auriculate, sometimes wanting, limb entire or bifid;

coronal scales present. Stamens: filaments glabrous

or barbellate in proximal half Ovary partially septate.

Capsule oblong or ovoid-oblong. Seeds reniform, flanks

excavate, back flat or furrowed, radially colliculate-

echinulate.

± 20 spp. Circum-Mediterranean, extending to

Macronesia and Pakistan; introduced elsewhere.

10. Silene nocturna L., Species plantarum 1: 416

(1753). Type: ‘Habitat in Italia, Pensylvania’, Herb.

Linn. 583.8 [LINN, lecto. — digital image!, designated

by Ghafoor: 91 (1978)].

Annual to 50 cm tall; stems erect, wand-like, sim-

ple or branched, 0.75-1.50 mm diam., lower parts

closely puberulous with very short, spreading hairs

mixed with longer hairs up to 1 mm long; upper parts

of stems and inflorescences closely puberulous with

very short, spreading hairs mixed with short, spreading,

gland-tipped hairs. Leaves mostly basal, oblanceolate

to spathulate-oblanceolate becoming oblong-lanceolate

distally, 15-50 x 5-15 mm, apiculate or obtuse, base

narrowed in lower leaves, ± densely pubescent with

± appressed hairs, mixed on upper leaves with gland-

tipped hairs, margins towards base ciliate with hairs to

1 mm long, without evident side veins. Inflorescence a

secund, spike-like monochasium terminating main stems

and branches, (3-)5-8-flowered. lower flowers remote;

bracts smaller than leaves, unequal, fertile bract linear to

awl-shaped, densely puberulous with gland-tipped hairs

mixed with acute, eglandular hairs; pedicels up to 12

mm long in lower flowers and elongating to 35 mm long

in fruit, mostly 3-4 mm long in upper flowers or more

rarely upper flowers subsessile, lowermost pedicels

sometimes patent or deflexed in fruit. Calyx narrowly

um-shaped in flower, 9-12 x 2. 0-2. 5 mm at anthesis,

equally 10-veined, reticulately veined in distal half,

densely adpressed-puberulous with slightly longer hairs

0.5 mm long on veins, lobes narrowly triangular-lanceo-

late, ±2 mm long, densely ciliolate. Flowers suberect at

anthesis, unscented, nocturnal or cleistogamous. Petals

pink, claw 6-8 mm long, oblong but narrowed below,

without auricles, limb bifid or emarginate, 1.5^.0 x

1.5^.0 mm, coronal scales 0.25-0.50 mm long. Sta-

men filaments unequal, 5-7 mm long, shorter series

shortly included, longer series shortly exserted. Ovary

pyriform, ± 4 mm long; styles ± 2 mm long, deeply

included. Capsule ovoid, 6-8 x 4 mm, ± 4-5 x longer

than carpophore, minutely transversely rugulose; carpo-

phore ± 1. 5-2.0 mm long, pubescent. Seeds 0. 5-1.0 mm,

reniform with hilar notch, reddish brown, face excavate-

auriculate, testal cells radially elongated-fusiform, col-

liculate, back broad with narrow, tuberculate groove.

Flowering time: (Jun.-)Aug.-Nov. Figures 1C, D, 19.

Vernacular name: Mediterranean catchfly.

Ecology and distribution: widespread through the

Mediterranean, extending to Portugal and Northern

Spain; introduced and established in western Australia

and an occasional introduction in the southwestern

Cape, where it has been recorded mainly from the Cape

Peninsula but also from Bredasdorp, with a recent record

from Sardinia Bay west of Port Elizabeth (Figure 20).

The species appears to be restricted to disturbed ground

and roadsides.

The species was first collected in South Africa under

the name Silene Ipendula L. by Carl Thunberg (Table

1), who travelled in the country from 1772 to 1774.

Another early South African record of Silene nocturna

that we have seen was made by Carl Zeyher {Zeyher

1960) in the mid-nineteenth century, somewhere in the

vast tract of country between Malabarshoogde [near

182

Bothalia 42,2 (2012)

FIGURE 19. — Silene noctwna, Sardinia Bay, Goldhialt & Porter-

13661 . A, flowering stem; B, calyx laid out; C, petal; D, gyn-

occium; E, gynoecium 1/s and t/s; F, seed side view; G, seed dor-

sal view. Scale bar: A, 1 0 mm; B, 5 mm; C, D, 2.5 mm; E, 2 mm;

F, G, 0.5 mm. Artist: John Manning.

Queenstown in Eastern Cape] and Hessaquaskloof [near

Riviersonderend in Western Cape]. The next collection,

from Wynberg Hill, dates from the end of the nineteenth

century. Certainly, within two decades of this latter col-

lection, the species was being found at several other

localities around the Peninsula but has so far failed to

establish itself as invasive. These early collections were

FIGURE 20. — Recorded southern African distribution of Silene gallica,

• ; S. noctwna, o.

misidentified as either S. gallica or S. clandestina [now

S. aethiopica] or were left unnamed.

Diagnosis and relationships: the occurrence of Silene

nocturna in southern Africa has been overlooked until

now due to confusion with S. gallica, another introduced

annual. The two species are superficially similar veg-

etatively and can be confused as herbarium specimens,

although S. nocturna usually has more slender and lax

flowering stems, typically with the lowermost flowers on

long pedicels up to 35 mm long in fruit. They are best

distinguished by details of stem vestiture, and especially

by their flowers and seeds. Both species have a mixture

of short and long eglandular hairs on the lower portions

of the stem but in S. gallica the longer hairs are dis-

tinctly shaggy and 1-2 mm long, whereas in S. nocturna

they are shorter and at most up to 1 mm long. The flow-

ers of S. nocturna, as the name suggests, are typically

nocturnal with distinctly bilobed petals (Figure 19C),

and the calyx is appressed-puberulous, without gland-

tipped hairs and with only slightly longer hairs on the

veins (Figure 19B). S. gallica, in contrast, has diurnal

flowers with distinctive, unlobed petals (Figure 2 1C),

and a glandular-haired calyx with characteristic glassy

bristles 2-3 mm long on the veins (Figure 2 IB). Both

species have seeds with deeply excavated flanks but dif-

fer in detail: seeds of S. nocturna have spreading, almost

wing-like shoulders with a narrrowly grooved back (Fig-

ure 1C, D, 19F, G), and those of S. gallica have rounded,

unwinged shoulders and a flat or slightly concave back

(Figure IE, F, 21G, H).

Additional specimens

WESTERN CAPE. — 3318 (Cape Town): mountain side above

Clifton, (-CD), Aug. 1913, Kensit s.n. (BOL); Cecilia boundary, (-

CD), 16 Sept. 1945, Levyns 17426 (BOL); Rondebosch, (-CD), Sept.

1902, Anon STE 13264 (NBG); Cape Peninsula, between Bishopscourt

and Kirstenbosch, (-CD), 3 Oct. 1951, Salter 9048 (BOL); Bishop-

scourt, (-CD), Nov. 1951, Pillans 10646 (MO); Paradise Estate, N

slopes of Wynberg Hill, (-CD), Sept. 1918, L. Bolus s.n. (MO); Suider

Paarl, (-DB), Aug. 1917, Roberts & Adendorf 17671 (PRE); Tyger-

berg Nature Reserve, (-DC), 6 Sept. 1975, Loubser 3380 (NBG);

Sept. /Oct. 1976, Loubser 3384 (PRE). 3418 (Simonstown): Wyn-

berg Hill, (-AB), 8 Nov. 1896, Wolley Dod 1817 (BOL); E slopes of

Vlaggenberg [Vlakkenberg], 19 Sept. 1915, Pillans 2832 (BOL). 3420

Bothalia 42,2 (2012)

FIGURE 2 1 . — Silene gallica, Kleinmond, without voucher. A, flower-

ing plant; B, caljot laid out; C, petal; D, petal limb [var. quin-

quevulnera]', E, androecium laid out; F, gynoecium; G, seed side

view; H, seed dorsal view. Scale bar: A, 10 mm; B, 5 mm; C-F,

2.5 mm; G, H, 0.5 mm. Artist: John Manning.

183

(Bredasdorp): De Hoop, flats in Grootwiidkamp, (-AD), 2 Aug. 1979,

Burgers 2144 (NBG).

EASTERN CAPE. — 3425 (Skoenmakerskop): Sardinia Bay, road

side at Loerie road intersection, (-BA), 18 Sept. 2010, Goldblatt &

Porter 13661 (MO, NBG). Locality uncertain'. South Africa, between

Malabarshoogde and Hessaquaskloof, Sept, [without year], Zeyher

1960 SI 1-24 179 [left hand specimen] (S).

11. Silene gallica L., Species plantarum: 417 (1753),

nom cons.; Sond.: 127 (1860). Type: ‘Habitat in Gallia’,

Herb. Linn. 583.11 [LINN, lecto.— digital image!, desig-

nated by Greuter: 102 (1995a)].

Silene anglica L.: 416 (1753), nom. rej. in favour of

S. gallica L. S. gallica var. anglica (L.) W.D.J.Koch: 109

(1843). Type: ‘Habitat in Anglia, Gallia’, Herb. Linn.

583.1 [LINN, lecto., designated by Talavera & Munoz

Garmendia: 498 (1989)].

[see Chater & Walter (1964) and Greuter (1997) for

additional heterotypic synonyms].

Annual to 40 cm tall; stems ascending or erect, sim-

ple or well-branched from base and above, 0.75-3.00

mm diam., lower parts closely puberulous with very

short, ± deflexed or spreading hairs mixed with longer,

shaggy, spreading or upcurved hairs 1-2 mm long but

later glabrescent; upper parts of stems and inflorescences

closely puberulous with very short, ± deflexed or spread-

ing hairs mixed with short, spreading, gland-tipped

hairs, inflorescence axis especially densely gladular-

pubescent. Leaves oblanceolate to spathulate-oblanceo-

late becoming oblong-lanceolate distally, 15-50 x 4-15

mm, apiculate, base narrowed in lower leaves, sparsely

or densely pubescent with erect or ± appressed hairs,

mixed on upper leaves with gland-tipped hairs, mar-

gins towards base ciliate with long, straggling hairs to

3 mm long, without evident side veins. Inflorescence a

secund, spike-like monochasium terminating main stems

and branches, (4-)8-15(-20)-flowered, lower flowers

often remote; bracts smaller than leaves, unequal, fer-

tile bract linear, sterile bract larger and more leaf-like,

densely puberulous with gland-tipped hairs mixed with

acute, eglandular hairs; pedicels mostly up to 3 mm long

in lower (rarely up to 12 mm long in lowermost flower

developed in axil of upper most leaf) but upper flowers

subsessile. Calyx sub-cylindrical in flower, 8-10 x 1.5-

2.0 mm at anthesis, equally 10-veined without reticulate

venation, densely puberulous with mix of gland-tipped

and acute hairs and with conspicuous, glassy bristles

2-3 mm long on veins, lobes awl-like, 2-3 mm long.

Flowers suberect at anthesis, unscented, diurnal. Pet-

als longitudinally twisted and propellar-like, white or

pink, sometimes with large crimson blotch or stain, claw

6-8 mm long, oblong, without auricles, limb obovate

or oblong, minutely crenulate, unlobed, 1.5^.0 x i.O-

2.5 mm, coronal scales ± 1 mm long. Stamen filaments

unequal, shorter series ± 4-6 mm long, included, longer

series ± 7-9 mm long, exserted up to 2 mm, puberulous.

Ovary pyriform, ± 2 mm long; styles ± 1.5 mm long,

deeply included. Capsule ovoid, 6-8 x 4-5 mm, ± 6-8 x

longer than carpophore, minutely transversely rugulose;

carpophore ± 1 mm long, pubescent. Seeds 0. 7-1.0 mm,

reniform, greyish brown, face concave-excavate, testal

cells radially elongated-fusiform and mostly 2-4 granu-

late, back flat or weakly concave, tuberculate. Flowering

time: (Jun.-)Aug.-Nov. in the southwestern and south-

184

Bothalia 42,2 (2012)

em Cape; Sept.-April in the summer-rainfall region.

Figure IE, F, 21.

Vernacular names: small-flowered catchfly, gunpow-

der weed (Sonder 1860), joppies (Hanekom 1005), hard-

ebolkeiltjies [hardebolletjies] (Wagner s.n. STE16912).

Distribution and ecology: native through Europe

and western Asia; now introduced as a weed into many

parts of the world, and well established in South Africa,

mainly in the winter-rainfall region, where it is most

common in the southwestern Cape, ranging northwards

onto the Bokkeveld Escarpment and the Kamiesberg

in central Namaqualand, and eastwards along the East-

ern Cape coast to Grahamstown and East London, with

isolated records further north, in southern and central

KwaZulu-Natal and Potchefstroom (Figure 20); it is also

recorded from the the highlands of east tropical Africa

in Kenya, Tanzania, and Uganda (Turrill 1956a). S. gal-

lica typically occurs as a weed of agricultural lands and

in waste places, along roadsides, and in other disturbed

sites, but in some places has also invaded more open

native vegetation that has been subjected to light distur-

bance, such as trampling and grazing.

The species was already established in South Africa

by the late eighteenth century, when it was encountered

here by Carl Thunberg, who travelled in the country

from 1772 to 1774. Another early collection made near

Port Elizabeth by Ecklon in the 1830s was misidentified

as S. cernua Thunb., a later synonym of S. aethiopica

Burm.

Diagnosis and relationships: an erect or somewhat

spreading, rarely ±prostrate annual with obovate-spathu-

late lower leaves, densely glandular-puberulous on the

upper parts of the stem and on the inflorescence, and

highly characteristic flowers with ± entire petals. The

calyx is ± tubular at anthesis with distinctive glassy

bristles 2-3 mm long on the veins, and the um-shaped

capsule, 6-8 mm long, is carried on a very short carpo-

phore, ± 1 mm long. The styles are very short, ± 1.5 mm

long.

The seed testal cells are radially elongated on the

flanks, which are deeply excavated, and each cell is

ornamented with a single series of 2-4 granules (Figure

IE). A double series of cells along the midline demar-

cates the flat or weakly convex seed back (Figure IF).

Minor variants have been distinguished in the past,

based on branching, flower colour, petal shape, and dif-

ferences in capsule orientation (Greutcr 1995a) but only

the striking colour form, var. quinquevulnera, is recog-

nized here. This variant cannot be distinguished from

the typical form on any other grounds and identifica-

tion of herbarium specimens depends on colour notes

on the collecting labels or on residual coloration of the

petals. Both varieties are sometimes represented on a

single herbarium sheet. A second variety, var. anglica,

was listed by Sonder (1860) as applying to more highly

branched and spreading plants but as far as we can tell

this is purely a manifestation of growing conditions and

has no taxonomic value at all.

11a. var. gallica

Petals uniformly white to pink. Figure 2 1C.

Distribution: as for species (Figure 20).

lib. var. quinquevulnera (L.) WDJ.Koch, Synop-

sis florae germanicae et helveticae, ed. 2: 109 (1843);

Sond.: 127 (1860). Silene quinquevulnera L.: 416

(1753). Type: ‘In Lusatia, Italia, Gallia’, Herb. Purser

XI: 72 [UPS, lecto., designated by Talavera & Munoz

Garmendia: 409 (1989)].

Petals with large crimson spot or staining in centre of

petal limb. Figure 2 ID.

Distribution: this variant occurs mainly in the south-

western Cape, extending into the Eastern Cape as far as

Grahamstown (Figure 20), but has not been recorded

further east or north in southern Africa, nor from tropical

Africa (Turrill 1956a; Wild 1961). It occurs mixed with

the typical variety, usually in the minority.

Additional specimens

*Collections comprising or including individuals of

var. quinquevulnera.

NORTH-WEST.— 2627 (Potchefstroom): Potchefstroom, (-AC),

10 Oct. 1903, Burn Davy 1761 (NH).

KWA-ZULU NATAL. — 2930 (Pietemaritzburg): Lidgetton, (-

AC), 1 April 1917, Mogg 624 (BOL); Lions River Dist., Umgeni

above Midmar, (-AC), 13 Oct. '964, Moll 1246 (NU); Pietermaritz-

burg, (-CB), Sept. 1946, Huntley 51 (NH, NU); 1968 [without month],

Garrett 76 (NU); Inchanga, (-DA), Sept. 1955, Alexander 14 (NU);

Umlaas Road, (-DA), 4 Sept. 1981, Manning 18 (NU); Botha’s Hill,

(-DC), 30 Sept. 2003, Styles 1597 (NU). 2931 (Stanger): Durban flat,

(-CC), Sept. 1883, Wood 2245 (NH). 3029 (Kokstad): Harding, Farm

Bedford, (-DB), 28 Sept. 1963, Lennox s.n. (NU).3030 (Port Shep-

stone): St Michaels-on-Sea, (-CD), 10 Oct. 1973, Mogg 38284 (NH).

3130 (Port Edward): Port Edward, (-AA), without date or collector,

NU2490/3 (NU). Imprecise locality: Natal, received Feb. 1884, Wood

1932 (BOL).

NORTHERN CAPE, — 3017 (Hondklipbaai): Kamiesberg, Skilpad

Wildflower Reserve, (-BB), 29 Sept. 1995, Cruz 120 (NBG). 3119

(Calvinia): top of Vanrhyns Pass, (-AC), 1 Oct. 1947, Compton 2867

(NBG); Oorlogskloof Nature Reserve, (-CA), 16 Oct. 1996, Pretorius

394 (NBG).

WESTERN CAPE.— 3118 (Vanrhynsdorp): Gifberg, (-CD), 16

Sept. 1911, Phillips 2490 (NBG). 3218 (Clanwilliam): Clanwilliam,

Olifants River Dam, (-BB), 10 Sept. 1949, Barker 5737 (BOL, NBG).

3219 (Wuppertal): Cold Bokkeveld, Ondertuin, (-CC), 20 Dec. 1980,

Hanekom 2610 (PRE); Cold Bokkeveld, Skoongesig, (-DC), 4 Nov.

1967, Hanekom 1005 (NBG, PRE). 3318 (Cape Town): Signal Hill,

(-CD), Sept. 1887, Thode 9236 (NBG); Cape Town, slopes above

De Waal Drive, (-CD), 26 Oct. 1928, Gillett 1819* (NBG, PRE);

slopes of Table Mt above Cape Town, (-CD), 26 Oct. 1928, Hutch-

inson 997* (BOL); E slopes of Lions Head, (-CD), Sept. 1913, Ken-

sit s.n. BOL45322 (BOL); Raapenberg, (-CD), Dec. 1890, Guthrie

840 (BOL); near Rondebosch, (-CD), 8 Sept. 1895, Wolley Dod 96*

(BOL); Rondebosch, (-CD), 7 Aug. 1938, Adamson 1926 (BOL);

Kuils River, Langverwacht, (-DC), 26 Oct. 1971, Oliver 3708 (NBG);

Jonkershoek, Bosboukloof, (-DC), Oct. 1967, Kerfoot 6080 (PRE);

Bonterivier Farm, SW of Stellenbosch, (-DD), 6 Oct. 1989, Buys 56*

(NBG); Stellenbosch, Voeitjiesdorp, (-DD), 10 Sept. 1978, Boucher

3928* (NBG); Stellenbosch, (-DD), 19 Sept. 1966, Taylor 6888*

(PRE, NBG). 3319 (Worcester): Ceres, Bokkerivier Farms, (-AD),

19 Nov. 1963, Booysen 95 (NBG); Worcester High School, (-CB),

18 July 1980, Walters 2078 (NBG); Worcester, Langerug Koppie, (-

CB), 24 Aug. 1977, Walters 1880 (NBG). 3320 (Montagu): Cogmans

Kloof, (-CC), 18 Oct. 1964, Bayliss 2429 (NBG). 3418 (Simonstown):

Fishhoek, (-AB), July 1918, Pahe s.n. BOL45321 (BOL); Noordhoek,

( AB), Sept. 1937, Eames s.n. BOL45325 (BOL); Kommetjie vlei,

Farm Imhoffs Gift, (-AB), Davies 21 (NBG, PRE); 5 km from Hout

Bothalia 42,2 (2012)

185

Bay, (-AB), 14 Oct. 1980, Davies 40* (NBG); Faure, (-BA), 14 Sept.

1946, Jordaan s.n. STE2490* (NBG); Somerset West, (-BA), 10 Aug.

1944, Parker 3898* (BOL, NBG); Betty’s Bay, Harold Porter Botanic

Reserve, (-BD), 11 Sept. 1968, Ebersohn 52/68* (NBG); Kogelberg

State Forest, along road near 2"‘‘ dwelling, (-BD), 29 Nov. 1991,

Kruger 102 (NBG). 3419 (Caledon): Caledon, Zwartberg [Swartberg],

(-AB), 30 Sept. 1980, Hilliard & Biirtt 13068 (NU); Palmietrivier near

Kleinmond, (-AC), Sept. 1930, Rossouw s.n. NBG 1 1240 (NBG); ± 3

km from Greyton on Genadendal Road, (-BA), 29 Sept. 1997, Meyer

1402* (PRE); 5 miles [8 km] NW of Riviersonderend, (~BB), 18 Sept.

1949, Heginbotham 100 (BOL, NBG); Napier, (-BC), Jan, 1937 [fruit-

ing], Jordaan 834 (NBG); Caledon/Napier road ± 1 km E of Stan-

ford/Rietpoel crossroads, (-BD), 25 Aug. 1995, Palerson-Jones 565

(NBG); Fairfield Farm, (-BD), 3 Oct. 1994, Kemper IPC657 (NBG);

Groot Hagelkraal, (-DA/DC), 25 July 1995, Paterson-Jones 470

(NBG); Bredasdorp, Anyskop, (-DD), Jan. 1937 [fruiting], Jordaan

137 (NBG). 3420 (Bredasdorp): Swellendam, (-AB), 25 Aug. 1956,

Theron 2052 (BOL); Potberg, (-BC), 20 Aug. 1980, Burgers 2484

(NBG).

EASTERN CAPE.— 3226 (Fort Beaufort): Cold Springs, (-BA), 9

Oct. 1983, Jacot Guillarmon 9245 (PRE); Grahamstown. (-BC), Oct.

1888, Galpin 227 (PRE); Grahamstown Memorial Garden, (-BC), 9

Oct. 1988, Jacot Guillarmod 10120* (PRE); Port Alfred. (-DB), 28

Sept. 1918, Britten 806 (PRE); Fort Hare Farm, (-DD), 11 Aug. 1978,

Gibbs Russell 4382 (PRE). 3227 (Stutterheim): Albany, (-AC), 6 Nov.

1965, Bayliss 2998 (NBG); Albany Dist., Howison’s Poort, (-AC),

16 Oct. 1966, Bayliss 3645 (NBG); Fort Cunnyngham, (-AC), Sept.

1897, Sim 968 (NU), 2181 (PRE); cultivated ground near Komgha,

(-BD), Nov. 1891, Flanagan 1181* (BOL, PRE); Grahamstown, Set-

tler’s Monument, (-BC), 5 Oct. 2004, Ramdhani 496 (NH); Stutter-

heim Dist., Kologh State Forest, (-CB), 7 Dec. 2001, Klein 78* (PRE);

King Williamstown, (-CD), Nov. 1891, Sim 971 (NU); East London,

(-DD), Oct. 1963, Bokelmann 8 (NBG). 3228 (Butterworth): Kentani,

(-CB), 19 Oct. 1906, Pegler 1412 (PRE). 3322 (Oudtshoom): Bas-

sonsrus, upper Cango Valley, (-AC), 4 Nov. 1974, Moffett 434 (NBG).

3323 (Willowmore): near Avontuur, (-CA), Nov. [without year], Bolus

2259 (BOL). 3325 (Port Elizabeth): Zuurberg Mountains, (-BC), 10

Oct. 1975, Bayliss 7114* (NBG); Uitenhage Dist., sandy hills near

the Zwartkop River, (-DC), Aug.-Oct. [without year], Ecklon 431

(BOL, NBG); Port Elizabeth, (-DC), July 1942, Cruden 448 (BOL);

15 Sept. 1982, Immelman 353 (PRE); Red House, (-DC), Sept. 1914,

Paterson 16232 (PRE). 3421 (Riversdale): Reisiesbaan siding, (-AB),

28 Aug. 1979, Bohnen 6351 (NBG). 3422 (Mossel Bay): road from

Kleinbrakrivier to Gannakraal along Moordkuilrivier valley, (-AA), 19

Oct. 1990, Joffe 878 (NBG); Pinedew farm, E of Wilderness, (-BA),

7 Nov. 1979, Hugo 1926 (NBG). 3423 (Knysna): Knysna, Belvedere,

(-AA), 9 June 1921, Duthie s.n. STE29219 (NBG); Plettenberg Bay,

(-AB), Sept. 1921, Smart sub Rogers 15459 (PRE); Ratels Bosch, (-

BA), June 1908, Fourcade 282 (BOL). 3424 (Humansdorp): Witte Els

Bosch [Witelsbos], (-AA), Nov. 1922, Fourcade 2411* (BOL, NBG);

Humansdorp, (-BB), June 1932, Wagner s.n. STE16912 (NBG);

Humansdorp Dist., Modderfontein, (-BB), 19 Feb. 1932 [cult. Onder-

sterpoort], Steyn s.n. (PRE); Jeffrey’s Bay, (-BB), 17 Dec. 1956, Tay-

lor 5154 (NBG).

ACKNOWLEDGMENTS

The following colleagues kindly located type speci-

mens in their collections and provided us with digital

images: Nicolas Fumeaux of the Conservatoire et Jardin

botaniques de la Ville de Geneve (G); Jens Klacken-

berg (Curator) and Mia Ehn of the Swedish Museum of

Natural Flistory (S); and John Hunnex from the British

Museum of Natural History (BM). Thanks to Dr. Bruno

Wallnofer, Curator, Vascular Plant Collections, Naturhis-

torisches Museum, Vienna (W), for confirming the loss

of the type of Silene caffra Fenzl ex C.MuelL; Martin

Callmander, Missouri Botanical Garden for assistance

with types; Roy Gereau, Missouri Botanical Garden,

for advice on nomenclatural matters; Mary Stiffler, Mis-

souri Botanical Garden, for help with literature searches;

Anne-Lise Fourie of the Mary Gunn Library for allow-

ing us access to the De Candolle Herbarium micro-

fiche; Esmerialda Klaassen of the Windoek Herbarium

(WIND) for providing some critical measurements; and

Miranda Waldron, SEM Unit, University of Cape Town

for technical assistance. Ingrid Nanni provided material

of Silene vulgaris for illustration, and Stephen Boat-

wright prepared the electronic figures. We are also grate-

ful to all curators for allowing us access to their collec-

tions, and to Bengt Oxelman for valuable comments on

the manuscript.

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Bothalia 42,2: 187-215 (2012)

Notes on African plants

VARIOUS AUTHORS

FABACEAE

OPHRESTIA OBLONGIFOLIA (PHASEOLEAE) IN SOUTHERN AFRICA

INTRODUCTION

Ophrestia H.M.L. Forbes is a member of the sub-

tribe Ophrestiinae in tribe Phaseoleae (Fabaceae). This

sub-tribe was erected by Lackey (1977) to include three

genera: the mostly African Ophrestia, the tropical Afri-

can Pseudoeriosema Hauman (both previously in Gly-

cininae) and the South-East Asian Cniddasia Prain (for-

merly in Galactiinae). Although the Ophrestiinae appear

to have affinities with some Asian Tephrosia Pers.

(Tephrosia is currently placed in the tribe Millettieae),

Lackey (1977) decided to keep them within the Phase-

oleae due to their twining habit. The Ophrestiinae can

therefore be described as "Tephrosia-\\k.Q Phaseoleae’.

A cladogram generated from a combination of molecu-

lar datasets (including Kajita et al. 2001 — rbcL\ Hu et

al. 2002 — ITS; and Wojciechowski et al. 2004 — matK)

nonetheless points to a sister relationship of Ophrestii-

nae with the core-Millettieae (Schrire 2005). These data-

sets, however, included only two representatives of the

Ophrestiinae, i.e. Ophrestia hedysaroides (Willd.) Verde,

and O. radicosa (A. Rich.) Verde.

Ophrestia, an anagram of Tephrosia, was cre-

ated by Forbes (1948) to accommodate T. oblongifolia

E. Mey. and three other species, which she described as

new at the time: O. nervosa H.M.L. Forbes, O. retusa

H.M.L. Forbes and O. swazica H.M.L. Forbes. The lat-

ter three were all subsequently placed into the synonymy

of O. oblongifolia (E.Mey.) H.M.L. Forbes by Verdcourt

(1970), who argued that the leaf characters given by

Forbes (1948) were “quite unusable and too variable”.

Ophrestia and Tephrosia (especially the Asian species)

share a number of characters, including pentafoliolate

leaves, silky petals and obscure stipels (Lackey 1981).

Ophrestia, however, differs from Tephrosia in having

leaflets with fewer, prominent and widely-spaced lateral

veins, which loop back without forming a marginal vein,

as opposed to the many, inconspicuous, and closely-

spaced parallel veins that form a marginal vein in Te-

phrosia (Forbes 1948; Verdcourt 1970).

Verdcourt (1970) expanded the circumscription

of Ophrestia by amalgamating it with Paraglycine

F. J.Herm. and Pseudoglycine F.J.Herm. on the basis that

Herman’s (1962) characters were not adequate to sepa-

rate these two genera from Ophrestia. Paraglycine is

characterized by having 1-7 leaflets, pubescent exterior

of corolla, and a swollen, cartilaginous, ariloid strophi-

ole (Herman 1962). Pseudoglycine has 5-7 leaflets,

sericeous standard on the outside towards the apex, and

a hilum covered by a membranaceous collar; it can also

be characterized by petaloid calyx lobes and a persist-

ent style in the fruit (Hermann, 1962). Ophrestia can

be characterized by lateral veins which loop back with-

out forming a marginal vein on the leaflets, it has 1-11

leaflets, pubescent corolla on the outer surface, and a

prominent aril. Some species of Ophrestia are superfi-

cially similar to certain species of Glycine Willd., which

would explain why a number of the species transferred

from Paraglycine fo Ophrestia were originally placed in

Glycine (e.g. Harms 1899; Baker 1929; Hauman 1955).

Ophrestia, however, differs from Glycine in having

short petioles, silky standard petals (glabrous in the lat-

ter), and prominent seed arils (Lackey 1977, 1981). In

addition. Glycine is generally trifoliolate, while Ophres-

tia is 1-5-jugate (Forbes 1948). As currently circum-

scribed (Schrire 2005), Ophrestia comprises 16 species

occurring mainly across Tropical Africa (8), and also

in Madagascar (4), Asia (3), and thinly distributed in

southern Africa (1). We describe and illustrate the south-

ern African O. oblongifolia for the first time since its

description, correct the typification, and also provide a

distribution map.

MATERIALS AND METHODS

Plant material was studied mainly from herbarium

specimens loaned from PRE, which has a comprehen-

sive collection of Ophrestia specimens. Habitat affinities

are described according to Mucina & Rutherford (2006).

Type specimens were viewed at www.plants.jstor.org.

TAXONOMIC TREATMENT

Ophrestia H.M.L. Forbes in Bothalia 4: 1003 (1948);

Verde. 24: 257 (1970); J.B. Gillett et al: 670 (1971).

Type species: O. oblongifolia (E.Mey.) H.M.L. Forbes

Paraglycine F.J.Herm.: 52 (1962); Hutch.: 448

(1964). Type species: P. hedysaroides (Willd.) F.J.Herm.

(= Ophrestia hedysaroides (Willd.) Verde.)

Pseudoglycine F.J.Herm.: 74 (1962); Hutch.: 448

(1964). Type species: P. lyallii (Benth.) F.J.Herm (=

Ophrestia lyallii (Benth.) Verde.)

Prostrate, climbing or erect perennial herbs or

shrubs, arising from a woody rootstock. Leaves digi-

tately or pinnately 3-1 1-foliolate or unifoliolate, leaf-

lets elliptic-oblong, ovate-oblong or lanceolate-oblong.

188

Bothalia 42,2 (2012)

with 5-7 prominent nerves above; stipules linear; sti-

pels minute or absent. Inflorescence axillary racemes,

slender, sometimes mueh longer than leaves, few- to

many-flowered. Bracts persistent, linear-lanceolate;

bracteoles persistent, linear or filiform. Calyx cam-

panulate, sometimes cylindric-campanulate, 5-lobed,

tube membranous; lobes subequal, all shorter than tube,

upper two ± connate. Corolla longer than calyx, purple,

purplish pink, violet, whitish pink or yellow; standard

oblong-pandurate, lower part auriculate, silky on outer

side, with well-developed, channelled claw; wings usu-

ally oblong, auriculate near base, with well-developed

linear claw, silky outside; keel narrowly elliptic, with

well-developed linear claw, auriculate near base, silky

outside. Stamens diadelphous, vexillary stamen free or

slightly joined to others, filaments alternatively long

and short, anthers all uniform in size. Ovary subsessile,

silky, ovules 2-8, style glabrous or hairy along one side,

stigma small, capitate. Fruit oblong or linear-oblong,

dehiscent, oblate, margins slightly thickened, glabrous

or slightly hairy, 2-5-seeded. Seeds oblong-ovate,

smooth, aril prominent.

16 spp.; southern Africa (South Africa and Swazi-

land), northwards into Tropical Africa, extending to Asia

and also in Madagascar.

Diagnostic characters'. Ophrestia shares a number

of characters with Tephrosia, notably the pentafolio-

late leaves, silky petals and obscure stipels, but is dis-

tinguished from it by the fewer, prominent and widely-

spaced lateral veins which loop back without forming a

marginal vein on the leaflets.

O, oblongifolia {E.Mey.) H.M.L. Forbes in Bothalia

4; 258 (1948); Verde. 24: 258 (1970). Tephrosia oblongi-

folia E.Mey.: 108 (1836); Meisn.: 86 (1843); C.Krauss:

54 (1846); Harv.: 209 (1862); O.Kuntze: 175 (1891);

Wood: 42 (1907); Burtt Davy: 377 (1932). Type: South

Africa, [KwaZulu-Natal], Drege s.n. P03453602 (P,

lecto.!, here designated). [Syntypes: [Eastern Cape],

‘ad Omsamcaba’ [Msikaba River], Drege s.n. (syn., not

located); ‘prope Omtendo et’ [Mtentu River], Drege

s.n. (P!, K!, syn.)]. [Note'. The fruiting specimen in P is

chosen as lectotype because it was annotated by Meyer

and because it is the only one with fruits (the fruits were

described in the prognosis)].

Glycinel wilmsii Harms: 302 (1899). Type: South

Africa, [Mpumalanga], “bei Lydenburg”, Wilms 383

(Bf, holo.; K, lecto.!, here designated).

O. nervosa H.M.L. Forbes: 1006 (1948). Type: South

Africa, [North-West], Marico district, Thode A 1394

(NH,holo.l).

O. retusa H.M.L. Forbes: 1005 (1948). Type: South

Africa, [North-West], N of Magaliesberg Dyer 8c Ver-

doorn 3405 (PRE, holo.!).

O. swazica H.M.L. Forbes: 1005 (1948). Type: Swa-

ziland, Dalriach at Forbes’ Reef //. Bolus 11845 (PRE,

holo.!; GRA!, K!, NH!, iso.).

Scandent perennial herb with trailing stems, up to 1

m wide. Leaves 1-5-jugate, rarely unifoliolate; leaf-

lets oblong, 40-75 x 20-30 mm, densely silky beneath,

glabrescent above, petiole (25-)45-80 mm long; stip-

ules linear-lanceolate, 5-7 x 1_2 mm; stipels absent.

Inflorescences few- to many-flowered axillary racemes;

peduncles 7-35 cm long. Flowers purple, purplish pink,

violet, whitish pink, yellow or bluish-purple, (6-) 11-1 5

mm long; bracts persistent, linear-lanceolate, 4-6 x ±

0.5 mm; bracteoles linear-lanceolate, 2-3 x ± 0.2 mm.

Calyx bilabiate, densely silky hairy, two upper lobes

fused up to two-thirds of their length, upper lip 6-8 mm

long, lower lip 6-8 mm long. Petals', standard oblong,

10-12 X 4-6 mm, silky on outside, claw 2-A mm long;

wings oblong, 6-10 x 4-6 mm, silky on outside, claw

± 3 mm long; keel narrowly elliptic, 5-9 x 3^ mm,

claw 3^ mm long. Stamens diadelphous, vexillary sta-

men slightly joined to others. Ovary 4-5 mm long, lin-

ear-oblong, 2-4-ovuled, style with few hairs along one

side, curved upwards. Fruit linear, laterally compressed,

20^0 X 7-8 mm, glabrescent, 1-2-seeded. Seeds brown

to black, oblong to ovoid, ±4 x ±3 mm, aril prominent.

Flowering time: Nov.-Feb. (Figure 1).

Distribution'. Ophrestia oblongifolia is the only spe-

cies of the genus that occurs in southern Africa and is

endemic to South Africa (Limpopo, North-West, Gau-

teng, Mpumalanga, and KwaZulu- Natal) and Swaziland

(Figure 2).

Diagnostic characters: close to O. hedysaroides and

O. radicosa in having generally oblong leaflets, however

in these two species the petioles are shorter than in O.

oblongifolia (the longest being up to 5.5 cm in O. radi-

cosa vs. 7.5 cm in the latter). In all African species of

Ophrestia (except O. hedysaroides), the leaflets are nar-

rower than in O. oblongifolia (up to 18 mm in O. uni-

foliolata (Bak.f ) Verde, compared to up to 2.5 cm). O.

oblongifolia also lacks stipels.

Key to varieties of O. oblongifolia'.

la. Leaves 3-7-foliolate, sparsely to densely covered with whit-

ish hairs . . . var. oblongifolia

lb. Leaves 3-11-foliolate; whole plant densely covered with

brownish hairs . . . var. velutinosa

O. oblongifolia var. oblongifolia

Leaves with 3-7 leaflets. Whole plant sparsely to

densely covered with whitish hairs.

Diagnostic characters'. Ophrestia oblongifolia var.

oblongifolia is less robust (with fewer leaflets than var.

velutinosa) and is sparsely to densely covered with whit-

ish hairs while in the latter the hairs are brownish.

Distribution and habitat: This variety is much more

widespread than var. velutinosa, occurring in South

Africa (Limpopo, North-West, Gauteng, Mpumalanga,

and KwaZulu-Natal) and the Mbabane area in Swazi-

land (Figure 2). It grows on sandy soil in the Savanna

and Grassland Biomes in the following vegetation types:

Zeerust Thomveld, Gold Reef Mountain Bushveld;

Central Sandy Bushveld, Makhado Sweet Bushveld;

Polokwane Plateau Bushveld, Legogote Sour Bush-

veld, Swaziland Sour Bushveld, Carletonville Dolomite

Grassland, KaNgwane Montane Grassland, KwaZulu-

Bothalia 42,2 (2012)

189

FIGURE 1. — Vegetative and reproductive morphology of Ophrestia oblongifolia (all from var. oblongifolia): A, flowering branch; B, flower in

lateral view; C, abaxial view of bract; D, lateral view of bracteole; E, calyx opened out with upper lobes to the left; F, standard petal; G, wing

petal; H, keel petal; I, stamens; J, pistil; K, lateral view of pod; L, lateral view of seed. Vouchers: A & B from S. Venter 11304 (PRE); C-J

from G.K. Theron 1311 (PRE); K, L, from B.J. Huntley 1551 (PRE). Scale bars: all 1 mm except K, 20 mm.

Natal Coastal Belt, Thukela Thomveld, and Ngongoni

Veld (Mucina & Rutherford, 2006).

Additional specimens

LIMPOPO. — 2329 (Polokwane): Polokwane, Hillside plot, Duven-

hageskraal Farm 686 LS, (-CD), 3 Dec. 1985, S. Venter 11304 (PRE).

2428 (Nylstroom): near Loubadspruit Bridge on road between Nyl-

stroom and Alma, (-CA), 4 Mar. 1985, M Welman 504 (PRE). 2429

(Zebediela): Percy Fyfe Nature Reserve (-AA), 22 Feb. 1968, B.J.

Huntley 1551 (PRE). 2430 (Pilgrim’s Rest): Graskop (-DB), 27 Nov.

1981, /l.-E. van Wyk 5204 (V'RE).

NORTH-WEST.— 2526 (Zeerust): Zeerust, (-CA), Mar. 1912,

T.J. Jenkins 11679 (PRE); Grasfontein, Lichtenburg, (-CC), 8 Dec.

FIGURE 2. — Distribution of Ophrestia oblongifolia var. oblongifolia,

• ; O. oblongifolia var. velutinosa, ■.

1929, J.D. Sutton 302 (PRE). 2527 (Rustenburg): Rustenburg Nature

Reserve, (-CA), 28 Dec. 1970, N. Jacobsen 1598 (PRE); Wolhuters

Kop (-DA), Dec. 1916, T. Nunns 9 (PRE). 2626 (Klerksdorp): NNE of

Lichtenburg, near side of the road to Roster, (-AA), 9 Mar. 1967, J.C.

Scheepers 1491 (PRE); 42 miles [68 km] from Roster on Lichtenburg

road, (-AB), 13 Oct. 1971, B. Clarke 249 (PRE); road to Ventersdorp,

22 km from turn-off of Roster/Lichtenburg road, (-BA), 8 Mar. 1985,

M. Welman 590 (PRE); 8 miles [13 km] NW of Ventersdorp, (-BD), 1

Feb. 1946, J.P.H. Acocks 12400 (PRE).

GAUTENG. — 2528 (Pretoria): camp adjoining govt house,

Bryntirion, (-CA), 14 Nov. 1926, C.A. Smith 3353 (PRE); Botanical

Reserve, Silverton, (-CB), 16 Dec. 1946, R. Story 1420 (PRE).

MPUMALANGA. — 2529 (Witbank): Loskopdam, Nooitgedacht,

(-AD), 12 Jan. 1967, G.K. Theron 1131 (PRE). 2531 (Romatipoort):

Rangwane, Songimvelo Game Reserve, Mlembe Mountain, (-CC), 7

Dec. 1992, G. Germishuizen 5683 (PRE).

SWAZILAND. — 2631 (Mbabane): Dalriach at Forbes Reef, (-

AA), H. Bolus 11845 (GRA, R, NH, PRE); Malolotja Nature Reserve,

(-AA), 18 Nov. 1985, L.M. Heath 418 (PRE).

RWAZULU-NATAL. — 2830 (Dundee): Hamewith, Mtunzini Dis-

trict, (-CB), 18 Nov. 1919, A.O.D. Mogg 5978 (PRE). 2930 (Pieterma-

ritzburg): Rey Ridge, (-DA), 18 Jan. 1987, P. Goldblatt & J. Manning

8395 (PRE); Inanda, Natal, (-DB), Jun. 1879, J.M. Medley-Wood 402

(PRE). 2931 (Stanger): Port Natal [Durban], (-CC), Drege s.n. c (P).

3030 (Port Shepstone): Mgayi, (-BC), 27 Jan. 1968, C.J. Ward 6351

(PRE); Uvongo Beach, (-CD), Mar. 1967, L.C.C. Liebenberg 7995

(PRE); Hibberdene, (-DA), 1 Jan. 1964, N. Grobbelaar 60 (PRE).

EASTERN CAPE. — 3129 (Port St. Johns): Lusikisiki, Umsikaba

River mouth, (-BC), 20 Jan. 1937, A.O.D. Mogg 13325 (PRE); Mkam-

bati Game Reserve, (-BD), 10 Dec. 1985, C Shackleton 368 (PRE).

3130 (Port Edward): Port Edward, (-AA), 5 Jan. 1981, G. Germishui-

zen 7727 (PRE).

O. oblongifolia var. velutinosa H.M.L. Forbes in

Bothalia 4: 1004 (1948). Type: South Africa, [KwaZulu-

190

Bothalia 42,2 (2012)

Natal], ‘Zululand’, Gerrard 1082 (NH, holo.!; BM!, K!,

P!, iso.).

Leaves with 3-11 leaflets. Whole plant densely cov-

ered with brownish hairs.

Diagnostic characters', this variety appears to be

more robust (with 3-1 1 leaflets as opposed to 3-7 leaf-

lets) than the typical variety, and much more hairy, with

brown rather than white hairs.

Distribution and habitat', var. velutinosa appears to be

restricted to the KwaZulu-Natal coastal belt, from Port

Shepstone to north of Durban. It occurs on sandy soil in

Midlands Mistbelt, Thukela Valley Bushveld and East-

ern Valley Bushveld (Mucina & Rutherford 2006).

Additional specimens seen

KWAZULU-NATAL.— 2931 (Stanger): Halfway between Tugela

Bridge and turn-off to Ultimatum Tree, (-AB), 14 Dec. 1972, N. Grob-

belaarl661 (PRE). 3030 (Port Shepstone): District Alexandria, Station

Dumisa, Farm Friedenau, (-AD), 5 Dec. 1908, H. Rudat'is 520 (PRE);

Dududu, Umkomaas, 2 km W of the Cedars Farm, (-BA), 5 Nov.

1992, A.M. Ngwenya 1051 (PRE); St. Michael-on-Sea, (-CB), 25 Dec.

1966, R.G. Strey 7090 (PRE).

ACKNOWLEDGEMENTS

We thank curator of PRE for assistance with herbar-

ium material, and the University of Johannesburg and

the National Research Foundation for financial support.

REFERENCES

BAKER, E.G. 1929. The Leguminosae of Tropical Africa I: 358. Eras-

mus Press, Gent.

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and Ferns of the Transvaal with Swaziland. South A frica: 418.

Longmans, Green & Co., London.

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Flora of tropical East Africa 3: 671. Royal Botanic Gardens,

Kew.

HARMS, H. 1899. Leguminosae africanae. 11. In A. Engler (ed.), Bota-

nische Jahrbiicher flir Systematik, Pflanzengeschichte und Pflan-

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(eds). Flora capensis 2: 239. Hodges, Smith, Dublin.

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Botanique de I’Etat a Bruxelles 25: 95.

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HU, J.-M., LAVIN, M., WOJCIESHOWSKI, M.F. & SANDERSON,

M.J. 2002. Phylogenetic analysis of nuclear ribosomal ITS/5.85

sequences in the Millettieae (Fabaceae): Poecilanthe-Cyclolo-

bium, the core Millettieae, and the Callerva group. Systematic

Botany 21: 122-157,.

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Press.

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J.J. 2001, rbcL and legume phylogeny, with particular reference

to Phaseoleae, Millettieae and allies. Systematic Botany 26:

515-536.

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und Natallandes. Regensburg, Germany.

KUNTZE, C.E.O. 1891. Revisio Generum Plantarum, Part 1: 175.

LACKEY, J.A. 1977. A revised classification of the tribe Phaseoleae

(Leguminosae: Papilionoideae), and its relation to canavanine

distribution. Botanical Journal of the Linnean Society 74: 163-

178.

LACKEY, J.A. 1981. Tribe 10. Phaseoleae. In: R.M. Polhill & P.H.

Raven, Advances in Legume Systematics 1: 301-327. Royal

Botanic Gardens, Kew.

MEISNER, C.D.F. 1843. Contributions towards a Flora of South Africa.

In: W.J. Hooker, The London Journal of Botany 2: 86. Hippolyte

Baillere, London.

MEYER, E.H.F. 1836. Commentariorum de plantes Africae australis 1:

65-75. Leopoldum Voss, Leipzig.

MUCINA, L. & RUTHERFORD, M.C. (eds). 2006. The vegetation of

South Africa, Lesotho and Swaziland. Strelitzia 19. South Afri-

can National Biodiversity Institute, Pretoria.

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kinder, & M. Lock, Legumes of the World: 393^30. Royal

Botanic Gardens, Kew.

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for the Flora of Tropical East Africa 11. Kew Bulletin 24: 257-

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WOJCIECHOWSKl, M.F., LAVIN, M. & SANDERSON, M.J. 2004.

A phylogeny of legumes (Leguminosae) based on analysis of

the plastid matK gene resolves many well-supported subclades

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A. N. MOTEETEE*t and B-E. VAN WYK*

’University of Johannesburg, Department of Botany and Plant Biotech-

nology, P.O. Box 524, Auckland Park 2006, Johannesburg, South Africa.

fCorresponding author: e-mail address: amoteetee(guj.ac.za.

MS. received: 2012-03-29

CYPERACEAE

IDENTITY AND TYPIFICATION OF CAREX COGNATA AND STATUS OF C. DRAKENSBERGENSIS

In a useful synopsis of sub-Saharan and Madagas-

can species of Carex, Gehrke (2011: 73) lectotypified

the name Carex cognat a Kunth on ‘South Africa, West-

ern Cape Province, Swellendam and George District,

W. Miindt s.n.' (S) (more correctly it was a neotypifica-

tion) and since that particular specimen is (in our opin-

ion) part of the same collection cited by Nees (1836) as

C. clavata Thunb. and currently identified by Gehrke

(2011) as such, the action resulted in Gehrke placing

C. cognata in the synonymy of C. clavata. Carex cog-

nata, in the sense of recent authors, e.g. Clarke (1898),

Kiikenthal (1909), Haines & Lye (1983), Gordon-Gray

(1995), and Verdcourt (2010) was treated by Gehrke

(2011: 75) as C. congolensis Turrill, citing several speci-

mens from southern Africa as this species. In the same

article, Gehrke (2011: 74) treated the evidently closely

related C. drakensbergensis C.B. Clarke as a separate

species. The aim of this brief note is to discuss and

clarify some of the issues of typification and synonymy

within this African species complex, mainly with refer-

ence to southern Africa.

Bothalia42,2 (2012)

191

Carex cognata Kunth

Kunth (1837) recognised that the name Carex re-

trorsa Nees (1836) was illegitimate and published C.

cognata as an avowed substitute. According to Art. 7.3

of the International Code of Botanical Nomenclature

(ICBN) (McNeill et al. 2006), the replacement name is

typified by the older, illegitimate name.

Until now, the identity of C. cognata has been in

little doubt. C. retrorsa Nees (1836: 204) was diag-

nosed against C. pseudocyperus L. by the accompany-

ing phrase ‘spiculis masculis geminis, femineis sub-

quatemis approximatis erectis cylindricis subsessilibus,

bracteis evaginatis longis, stigmatibus temis, fructibus

ovato-trigonis rostratis bifurcus striatis glabris squamam

lanceolatam setaceo-acuminatam serrulato-scabram

aequantibus, inferioribus reflexis’ [study of a large speci-

men sample has shown that in fact purely staminate

spikes vary between none and two and purely pistillate

spikes vary between two and six in number, while the

tubular sheathing portion of the lowermost inflorescence

bract varies from 0 to 20 mm long (Reid 1991)], and

this information was quoted verbatim by Kunth (1837)

when publishing his new name. Nees (and subsequently

Kunth) furthermore highlighted the similarity of his new

species to the very distinctive European plant C. pseii-

docypenis L. and Kiikenthal (1909) subsequently placed

it in the Section Pseudocypereae Tuck. It is difficult to

imagine that Nees and all reputable Cyperologists since

then could have misunderstood the affinities of this plant

and also could have confused it with the well-known and

(today still) quite common C. clavata Thunb., which is

described by Nees (1836) in detail in the same article

with specimen citations. During the course of her M.Sc.

study, Reid (1991) examined several of the collections

cited by Nees as C. clavata and verified the identifica-

tions, bearing in mind that all the original Carex mate-

rial seen by Nees had been lost (see further discussion

below). There is no doubt that the holotype of C. cog-

nata was still in existence and seen by Kiikenthal (1909:

699), who confirmed its identity while preparing his

monograph of Caricoideae.

Nees (1836: 205) cited the following specimen infor-

mation under C. retrorsa: Tn districtu “Zwellendam et

George” legit b. Mimdf (see Gunn & Codd (1981) for

the correct spelling of this name). In the same article

Nees cited two additional Mund collections, with the

same sparse information, under C. ecklonii (p. 203) and

C. clavata (p. 204) respectively. Our contention is that

Gehrke (2011) lectotypified (or more correctly neotypi-

fied) the name C. cognata on a sheet (acquired by S at

a much later date: see discussion below) of the Mund

collection that was known to Nees and cited by him as

C. clavata. Gehrke ’s (2011) synonymisation of C. cog-

nata under C. clavata was further motivated by her

belief that C. cognata, as understood by recent authors

e.g. Clarke (1898), Kiikenthal (1909), Haines & Lye

(1983), Gordon-Gray (1995) and Verdcourt (2010), did

not occur in the Western Cape, stating, under C. congo-

lensis, that “records from the Western Cape Province in

South Africa represent misapplications of the name”.

She apparently saw no coastal southern African records

at all, citing (as C. congolensis) only one specimen

from Lesotho (which Reid determined as depauperate

C. drakensbergensis C.B. Clarke) and one from Mpuma-

langa Province, South Africa. We, however, are of the

opinion that the species does occur in coastal southern

Africa, and cite below the relevant collections seen by

Reid (1991) for her M.Sc. study, plus one subsequent

record. The Mund locality is very imprecise, and in our

opinion it is plausible that Mund encountered this plant

somewhere in the ± 200 km between Swellendam and

George (probably closer to George: see Martin 4232

cited below), even though the population may no longer

be extant.

Article 9.17 of the International Code of Botanical

Nomenclature (ICBN) recommends that an erroneous

typification should be corrected once it becomes known.

The neotypification by Gehrke (2011 ) changes the appli-

cation of the name C. cognata and it is therefore relevant

to ensure that it is appropriate in accordance with Arts.

9.14 and 9.17 of the ICBN (McNeill et al. 2006).

It is firstly necessary to confirm that the Mund speci-

men on which C. retrorsa Nees was based has indeed

been lost. Mund’s specimens are at B, BR, PI, K, KIEL,

MO, and SAM (Gunn & Codd 1981); additionally, upon

his death in 1831 Mund’s personal herbarium went to

C.P. Ecklon (Gunn & Codd 1981) who distributed it on

behalf of Mund’s estate. Ecklon and Zeyher’s collections

were widely distributed, and amongst them are included

some Mund specimens, usually recognisable from the

label bearing some reference to the locality (Swellendam

and George District) and/or the collector (usually spelled

Mundt). In addition, Kunth’s personal herbarium of

70 000 specimens was acquired by B after his death in

1850. More relevantly to this issue, Nees’s herbarium of

± 10 000 specimens was acquired by B in 1855 (Hiepko

1987). Prior to this, Nees (1832, 1833, 1836) was evi-

dently studying Ecklon (and Zeyher)’s collections of

Cyperaceae and the titles of his publications suggest that

the Mund collections that he cited in these publications

were included with these specimens. With the exception

of types of Cyperns L. and Eleocharis R.Br. (in part),

nearly all Cyperaceae specimens were lost during World

War II (Pilger 1953). Ecklon himself died in 1868 and

his personal herbarium was in turn acquired by Sonder,

whose collections are now partly at S and partly at MEL

(Court 1972; Nordenstam 1980). All of the herbaria

mentioned above (Thiers, Index Herbariorum online)

were checked as far as possible but no material that

could conceivably be part of the original material relat-

ing to C. cognata Kunth was found.

Turning now to the diagnosis of C. retrorsa: ‘spiculis

masculis geminis, femineis subquatemis approxima-

tis erectis cylindricis subsessilibus, bracteis evagina-

tis longis, stigmatibus temis, fructibus ovato-trigonis

rostratis bifurcus striatis glabris squamam lanceolatam

setaceo-acuminatam sermlato-scabram aequantibus,

inferioribus reflexis. [Symbol for] Perennial’ translates

as ‘male spikes paired, females 4 or less, close together,

erect, cylindrical, sub-sessile, bracts non-sheath-

ing, long, stigmas 3, fmit ovate-trigonous, rostrate,

2-toothed, striate, glabrous, glumes [= spikelet bracts]

lanceolate, setaceous-acuminate, sermlate-scabrid,

equalling [in length the fmit], lowermost reflexed. Per-

ennial’. The glabrous utricle plus the spikelet bracts [±]

equalling the utricle in length, are in our opinion clearly

192

Bothalia 42,2 (2012)

diagnostic of C. cognata, but certainly not of C. clavata

in which the utricles are hairy, at least on the rostrum

margins, and the spikelet bracts are usually mueh shorter

than the utrieles.

C. drakensbergensis C.B. Clarke

Clarke (1898) distinguished C. drakensbergensis with

its longer, drooping, distant, pedunculate spikes with

dark ferruginous female bracts from C. cognata with

shorter, erect, clustered, nearly sessile spikes and green-

ish female bracts. Shortly afterwards Kiikenthal (1909)

found that the two taxa were not separable at species

level and reduced C. drakensbergensis to a variety of C

cognata.

Intensive herbarium studies of mainly southern Afri-

ean material (Reid 1991) indicated that plants matching

C. drakensbergensis occurred mainly at high altitude,

growing in open sunny habitats, whereas plants match-

ing C. cognata sensu stricto occurred near the eoast, in

the Okavango Swamps of Botswana, and [not reported

by Podlech (1967)] at waterholes in the Great Water-

berg of Namibia, growing in lightly shaded habitats. In

practice, however, it is not possible to divide herbarium

specimens into two meaningful taxa sinee there are

always intermediate examples. At Hogsback in Eastern

Cape, we observed individuals conforming to the con-

cept of C. drakensbergensis growing in open grassland

on a stream bank. A very short distanee downstream,

individuals of what is evidently the same population

were lightly shaded by planted Pinus species (the sedge

population apparently pre-dating the tree plantation) and

conformed to the concept of C. cognata. We concluded

that the differenees between these two taxa are entirely

habitat-related, and that C. drakensbergensis cannot be

upheld as a separate taxon, not even as a variety of C.

cognata, as was done by Kiikenthal (1909). Reid (1991,

unpublished) therefore regarded C. drakensbergensis as

conspecific with C. cognata, and Gordon-Gray (1995)

formally upheld this synonymy.

Gehrke (2011) once again treated C. drakensbergen-

sis as a separate species stating “Most South African

material so far identified as C. cognata can be assigned

to var. drakensbergensis, which I also regard as a dis-

tinct species (34. C. drakensbergensis). The remaining

material from South Africa and Lesotho usually has par-

tially pendulous spikes, which are less densely clustered

than most material from further north, and both forms

(i.e. those with clustered spikes and those with more dis-

tant ones) might deserve the status of subspecies. Carex

congolensis differs from 34. C. drakensbergensis by

spikes all erect or rarely with a pendulous lateral spike,

utricles inflated at maturity and a rostrellum with shorter,

1 mm-long, finer teeth (C. drakensbergensis has mostly

pendulous spikes, utricles not inflated at maturity and

a rostrellum with 1 mm long teeth).” However, having

studied numerous herbarium collections (Reid 1991), in

our opinion fully mature specimens of C. drakensber-

gensis do indeed have inflated utrieles, and as stated

above, the perceived differenees in orientation of spikes

and length of teeth on rostrellum (or rostrum) of the utri-

cles are infinitely variable within these taxa and cannot

be employed to separate them.

In light of the above arguments, the treatment of this

eomplex in southern Africa must be as follows:

C. cognata Knnth in Enumeratio Plantarum 2: 502

(1837), as a nom. nov. for C. retrorsa Nees {non sensu

Gehrke: 73 (2011) = C. clavata Thunb.]; Gordon-Gray:

39 (1995) [inch C drakensbergensis)', Verdeourt: 447

(2010) [as var. cognata). C. retrorsa Nees (1836), horn,

illegit. non Schweinitz (1824). C. pseudocyperus L.

var. cognata (Kunth) Boott: 141 (1867). Type: [West-

ern Cape] “In districtu ‘Zwellendam et George’ legit,

b. Mundt” (?|B, holo., not found). Neotype, designated

here: [Eastern Cape], ‘Distriet Kentani, along streams

where marshy, 1 200 ft.’ [± 365 m], 22 Sept. 1910, A.

Pegler 151 (PRE, neo.l). [The collections Pegler 151

from same locality, coll. 19 Sept. 1904 (BOL!), 19 Sept.

1907 (BOL!) and Nov. 1907 (GRA!) all represent Carex

clavata Thunb.].

C. drakensbergensis C.B. Clarke: 309 (1898); Gehrke:

74 (2011). C. cognata Kunth var. drakensbergensis

(C.B.Clarke) Kuk.: 699 (1909); Verdeourt: 447 (2010).

Type: East Griqualand [Eastern Cape], Kokstad, Vaal

Bank Farm, 18 Dec. 1889, W.J. Haygarth sub J.M.

Wood 4201 (Kl, lecto., designated by Gehrke: 74 (2011);

BOL!,NH!,isolecto.).

C. congolensis Turrill: 240 (1912); Gehrke: 75

(2011) [inch C. cognata Kunth]. C. cognata Kunth var.

congolensis (Turrill) Lye: 244 (1983); Haines & Lye:

384 (1983). Type: Congo [Democratic Republic of

Congo], Katanga, Elisabethville [Lubumbashi], 11°37'S,

27°24'E, 1 150 m, 21 Sept. 1911, Rogers 10082 (K,

leeto., designated by Gehrke: 75 (2011); BOL, isolecto.).

Carex cognata (inch C. drakensbergensis) appears

to be the only representative of Sect. Pseudocypereae

Tuck, in southern Africa. In common with most mem-

bers of the section, the leaves have a distinetive pattern

of cross-venation and together with the bright yellow-

ish-green leaf colour the species is quite distinctive and

easily recognised where it does oecur, even in the veg-

etative stage. The cross-venation is even more conspicu-

ous in dried material and easily observed with the naked

eye. Extensive fieldwork in southern Africa has shown

that the plants are rather sporadic in occurrence and

never form large populations. We think that the chief

means of long-range dispersal of these wetland plants

is by migratory aquatic birds bearing the fruit in mud

attached to their feet — hence the isolated oecurrence in

the Waterberg of Namibia. C. cognata sensu stricto also

occurs in the Okavango Swamps of Botswana and in

South Africa in Limpopo, Mpumalanga, and along the

coast of KwaZulu-Natal, and Eastern and Western Cape.

If C. congolensis with its synonyms is ineluded (see

Gehrke 2011), the speeies extends into Tropical Africa

as far north as Tanzania. Forms of the species previously

treated under the name C. drakensbergensis occur in the

midlands of the Eastern Cape, KwaZulu-Natal, and up

into the Drakensberg highlands from Eastern Cape and

Lesotho, extending through Mpumalanga and along the

eastern mountain chain onto the Vumba and Chimani-

mani Mountains of Zimbabwe and further north into

East Africa.

Bothalia 42,2 (2012)

193

Additional specimens (from coastal South A frica)

KWAZULU-NATAL. — 2732 (Ubombo): Ngwavuma District,

Kosi System, Sihadla, shade of tall hygrophilous trees on low bank

of stream in swamp forest, (-BB), 8 Jan. 1987, C.J. Ward 10050 (NH,

NU, PRE, UDW); Ubombo District, NE St. Lucia system, Pukwini,

marginal to swamp forest, (-DC), 9 Nov. 1972, C.J. Ward 8094 (NU,

PRE, UDW). 2832 (Mtubatuba): Hlabisa District, St. Lucia E shores

S of Tewate, in small clearing in swamp forest, (-AB), 16 Dec. 1964,

R.H. Taylor 422 (NH); Hlabisa District, Dukuduku East, light shade

just within margin of swamp forest, (-AD), 20 Nov. 1964, C.J. Ward

5080 (NH, NU, PRE, UDW); Lower Umfolozi District, Richards Bay,

common in patches in shade, swamp forest, (-CC), 28 Jan. 1949, C.J.

Ward 716 (NU, UDW); Lower Umfolozi District, Richards Bay, Sont-

wayo Pan, on margin, (-CC), 3 Feb. 1959, R.D. Guy & C.J. Ward 69

(NU, PRE). 2930 (Pietermaritzburg): Durban District, Isipingo Beach,

wet mud in reed swamp, (-DD), 4 Nov. 1950, C.J. Ward 1221 (NU,

UDW).

EASTERN CAPE. — 3129 (Port St. Johns): Port St. Johns, side of

ponds, roots in water, (-DA), Jan. 1929, H.A. Wager s.n. PRE39183

(PRE). 3228 (Butterworth); Transkei, Elliotdale, Cwebe Nature

Reserve, riverlet on N bank of Mbanyana River at bridge, (-BB), 22

Dec. 1992, E. Cloete 2304 (NH).

WESTERN CAPE. — 3418 (Simonstown): Wynberg District, Hout

Bay, near harbour, in very wet sand at foot of dripping cliff, (-AB), 2

Jan. 1972, RL. Forbes 421 (J); Simonstown District. Cape Peninsula,

Lakeside, at edge of the lake, (-AB), 10 Dec. 1939, M.R. Levyrts 7120

(B, BOL). 3422 (Knysna); Knysna District, [probably Groenvlei], (-

BB), ± 1960, A.R.H. Martin 4232 (K).

ACKNOWLEDGEMENTS

Thanks are due to Catherine Gallagher, Co-ordinator

of Curation, National Herbarium of Victoria, Australia,

who kindly provided images of Mund specimens of

Carex at MEL; also to all the herbarium curators who

kindly provided loans or images during the course of the

first author’s M.Sc. studies; and to Robert H. Archer for

critically reading this manuscript.

REFERENCES

BOOTT, F.M.B. 1 867. Illustrations of the genus Carex. Part 4: 127-234,

PI. 412-600. W. Pamplin, London.

CLARKE, C.B. 1898. Cyperaceae. In Thiselton-Dyer, W.T. (ed.). Flora

Capensis 7: 299-310. Lovell Reeve & Co., Ltd., London.

COURT, A.B. 1972. Preliminary notice on the Sender collection in the

National Herbarium of Victoria. Muelleria 2: 188.

GEHRKE. B. 2011. Synopsis of Carex (Cyperaceae) from sub-Saharan

Africa and Madagascar. Botanical Journal of the Linnean Soci-

ety 166: 5\-99.

GORDON-GRAY, K.D. 1995. Cyperaceae in Natal. Strelitzia 2. Natio-

nal Botanical Institute, Pretoria.

GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern

Africa. A. A. Balkema, Cape Town.

HAINES, R.W. & LYE, K.A. 1983. The sedges and rushes of East Afri-

ca. East African Natural History Society, Nairobi.

HIEPKO, P. 1987. The collections of the Botanical Museum Berlin-

Dahlem (B) and their history Englera 1: 219-252.

KUKENTHAL, G. 1909. Cyperaceae-Caricoideae. In Engler, A. (ed.).

Das Pflanzenreich 38: 1-824. W. Engelmann, Leipzig.

KUNTH, C.S. 1837. Enumeratio Plantarum 2. J.G. Cotta, Stuttgart &

Tubingen.

LYE, K.A. 1983. Studies in African Cyperaceae 27. Miscellaneous new

taxa and combinations. Nordic Journal of Botany 3: 241-244.

NEES AB ESENBECK, C.G. 1 832. Cyperaceae Capenses Ecklonianae.

Linnaea 7: 491-537.

NEES AB ESENBECK, C.G. 1833. Herm Professor Schraders Analecta

ad Floram Capensem verglichen mit Nees v. Esenbecks Erlaute-

rungen der Capschen Cyperaceen in Herm Ecklons Sammiung.

Linnaea 8: 75-94.

NEES AB ESENBECK, C.G. 1836. Cyperaceae Capenses Secundum

Novissimas Ecklonii Collectiones. Linnaea 10: 129-207.

MCNEILL. J., BARRIE, F.R., BURDET, H.M., BEMOULIN, V,

HAWKSWORTH, D.L., MARHOLD, K., NICOLSON, D.H.,

PRADO, J., SILVA, PC., SKOG, J.E., WIERSEMA, J.H. &

TURLAND, N.J. 2006. International Code of Botanical Nomen-

clature (Vienna Code), Electronic version, available at http://

ibot.sav.sk/icbn/main.htm.

NORDENSTAM, B. 1980. The herbaria of Lehmann and Sonder in

Stockholm, with special reference to the Ecklon and Zeyher col-

lection. Taxon 29: 279-291.

PILGER, R. 1953. II. Bericht uber den Botanischen Garten und Bota-

nische Museum zu Berlin-Dahlem, April 1947-Dezember 1948.

Willdenowia 1 : 22-3 1 .

PODLECH, D. 1967. Cyperaceae. In Merxmiiller, H. (ed.), Prodromus

einer flora von Sudwestafrika 165: 1-53.

REID, C. 1991. Systematics of the southern African species of Carex L.

(Cyperaceae). Unpublished M.Sc. dissertation. University of the

Witwatersrand, Johannesburg, South Africa.

SCHWEINITZ, L.D. 1824. An analytical table to facilitate the determi-

nation of the hitherto observed North American species of the

genus Carex. Annals of the Lyceum of Natural History of New

York \ 2: 65,-1 \.

THIERS, B. [continuously updated]. Index Herbariorum: A global

directory of public herbaria and associated staff. New York

Botanical Garden’s Virtual Herbarium, http://sweetgum.nybg.

org/ih/

TURRILL, W.B. 1912. Carex congolensis Turrill, in: Diagnoses Afri-

canae XLVIII. Bulletin of Miscellaneous Information. Royal

Botanic Gardens. Kew 1912: 240.

VERDCOURT, B. 2010. Carex L. In Hoenselaar, K., Verdcourt, B. &

Beentje, H.J. Cyperaceae. Flora of Tropical East Africa. Royal

Botanic Gardens, Kew.

C. ARCHER* and K. BALKWILL**

* National Herbarium, South African National Biodiversity Institute,

Private Bag XlOl, 0001 Pretoria, South Africa.

** Department of Animal, Plant, and Environmental Sciences, Univer-

sity of the Witwatersrand, Private Bag 3, 2050 Wits, Republic of South

Africa.

MS. received: 2011-09-12.

HYACINTHACEAE

ALBUCA TENUIFOLIA /cm A. 5'//Jfr7/(ORNITHOGALOIDEAE), TWO DISTINCT SPECIES FROM SOUTH AFRICA

INTRODUCTION

Albiica tenuifolia Baker was described and illus-

trated in 1 872 from South African material sent by Peter

MacOwan to William Wilson Saunders in England, nei-

ther citing a precise locality nor a herbarium collection

studied by Baker. This species has been mostly over-

looked and has only been cited in the recentmost check-

list of the South African flora (Manning & Goldblatt

2003), although with many uncertainties.

Recently, two very different taxonomic studies have

been published on Albiica tenuifolia Baker. On the one

hand Martinez- Azorin et al. (2011) presented data sup-

porting A. tenuifolia as a distinct species belonging to

A. subg. Mitrotepalum, after rediscovery of wild popu-

194

Bothalia 42,2 (2012)

lations fitting the concept described and illustrated by

Baker (1872). On the contrary, Muller-Doblies (2012),

almost at the same time, radically reinterpreted the cir-

cumscription of A. tenuifolia, treating the later A. shawii

as a synonym.

RECOVERY OF TWO DISTINCT SPECIES

The taxonomic proposal presented by Muller-Doblies

(2012) was based on a herbarium specimen collected

by Peter MacOwan (MacOwan 1851, GRA) (acronyms

of herbaria according to Thiers (2012)) and labelled as

“'Albuca temii folia Bak’. The existence of that herbarium

collection “allowed the taxonomic understanding of

Baker’s description”, although Muller-Doblies accepted

that “the voucher — MacOwan 1851, GRA — has no

similarity at all with the plate 335 in Saunders’ Refu-

gium”. Her explanation was that “the artist felt unsure of

the ‘wilting’ drooping flowers in this Albuca, as earlier

Albuca plates in the Refiigium Botanicum had mostly

erect flowers. Thus, the artist ‘corrected’ the drooping

flowers to erect ones and ‘corrected’ also somewhat the

size of flowers and pedicels. By ‘correcting’ the droop-

ing flowers to erect ones in a glandular species the art-

ist ‘created’ a new character combination not found in

nature, ‘a unique new species’.” She also assumes that

the description was written by Baker directly from the

illustration. Furthermore, she proposed that the fact

that MacOwan 1851 GRA has an identification label

with Baker’s handwriting, supposedly attached by Peter

MacOwan, supported its acceptance as material studied

by Baker, and hence she selected it as a ‘lectotype’. This

conclusion is not correct, however, since MacOwan’s

specimen is not original material (McNeill el al. 2006:

Art. 9.2 Note 2), and therefore should be regarded as a

neotype (McNeill et al, 2006: Art. 9.6 and 9.8). Muller-

Doblies (2012) speculated that a duplicate of MacOwan

1851 existed at Kew and was studied by Baker. This

could explain why Baker (1897) cited the same collec-

tion as A. minor. Even so, this fact does not change the

original concept of A. tenuifolia as described and illus-

trated in the protologue.

The taxonomic proposal made by Muller-Doblies

(2012) represents a radical change in the original concept

of A. tenuifolia, and it also modifies the taxonomy of a

widely accepted and common species in South Africa,

Albuca shawii. This compromises the correct application

of both names and here we clarify their taxonomy.

As argued by Martinez- Azorin et al. (2011), plants

fitting the original concept of A. tenuifolia were found

in several high altitude locations in the Sneeuberg, Great

Winterberg-Amatola, and Stormberg mountains, with an

outlying population in the Steenkampsberg in Mpuma-

langa. These plants are characterized by hypogeal and

proliferous, irregularly compressed bulbs; numerous,

filiform, minutely papillate (only visible under micro-

scope)— but not viscose glandulose — leaves; glabrous

peduncle and pedicels; erect, yellow-green flowers with

all six stamens bearing fertile anthers; a subglobose

ovary with divergent paraseptal crest; and a long, nar-

rowly obpyramidal trigonous style, fitting almost per-

fectly the protologue of Baker (1872). The erect flow-

ers, with six fertile stamens are characteristic of A. subg.

Mitrotepalum U.Mull.-Doblies (= A. sect. Branciona

(Salisb.) J.C. Manning & Goldblatt). There is, however,

an apparent incongruence regarding leaf indumentum in

Baker’s protologue. The leaves were described as ‘glan-

duloso-papillose under a lens’. This can be interpreted

as an error by Baker (1872), who likely interpreted the

papillae as glands. In any case, the structures are nei-

ther viscose nor stalked and evident to the naked eye, as

they are in A. shawii (see Figure 1 for comparison). This

fact is supported by the accurate illustration in the pro-

tologue of A. tenuifolia, which shows leaves, peduncle,

and pedicels glabrous at first sight, with a close-up of a

leaf section, numbered 1 in plate 335 of Baker (1872),

bearing minute papillae. Although the indumentum in

the illustration seems to be in part shortly stalked under

magnification this can be explained by drawing inaccu-

racies. In any case, the glands in A. shawii are always

relatively much longer with regard to leaf width. This

colour illustration was prepared by Walter H. Fitch, a

renowned artist who indeed reflected the character com-

bination found on living wild plants of this extant good

species. This plate was chosen as the obligate lectotype

FIGURE I. — Comparison of fresh leaf indumentum in A. tenuifolia

Baker (Eastern Cape, Somerset East, summit of Boscliberg,

Clark & Martinez-Azorln 23 GRA) and Albuca shawii Baker

(Eastern Cape, Linedrift, near Keiskamma river bridge on N2,

Martinez-Azorln, Dole! & Martlnez-Soler 520 GRA): A, Leaf

general view of A. tenuifolia (above) and A. shawii (below); B,

Sessile papillae of A. tenuifolia', C, Long stipitate glands oi A.

shawii.

BothaIia42,2 (2012)

195

of the species by Marti'nez-Azorin et al. (2011), who

also designated an epitype among specimens collected

on the summit of Boschberg (Somerset East).

The existence of the collection MacOwan 2071

(K000524291), that he identified as "A. polyphylla

Baker’, proves that MacOwan gathered plants from

the epitype locality. The sheet includes three complete

plants that were collected Tn proecipit. summi Mtis

Boschberg’, flowering in December at 4 500 feet alti-

tude, which fit perfectly the morphological traits, habi-

tat and locality of the epitype specimen of Albiica tenu-

ifolia. This collection was also cited as A. polyphylla by

Baker (1897: 459), probably because both taxa look very

similar when dried. However, the name A. polyphylla is

currently applied to a smaller, white flowered species,

characterized by the numerous filiform leaves spreading

falcately from the top of the bulb, and it occurs in drier

habitats at lower elevations.

It is also important to note that Baker (1873) in his

comprehensive revision of Omithogaleae and Chloro-

galeae added to description of his A. tenuifolia the fol-

lowing sentence: ‘Cap B. Spei, hort. Saunders, legit

MacOwan, v. v.’ Contrary to Miiller-Doblies (2012)

assumption, this demonstrates that Baker did see living

plants of A. tenuifolia grown in Saunders’s glasshouse,

of which no herbarium sheets were kept at K (see Baker

1897:461).

Regarding Albiica shawii, it was described by Baker

(1874) and is a species of A. subg. Falconera (Salisb.)

Baker (=.A. sect. Falconera (Salisb.) J.C. Manning &

Goldblatt) with nodding flowers, outer stamens usually

with sterile anthers, and leaves, peduncle and pedicels

with abundant and clearly visible, stipitate glands. Its

original description cited the following studied speci-

mens: "Caput Bona Spei ad ripas fluminis “Vaal river”

et in ditione Coles berg. Dr. Shaw!. Kaffraria in grami-

nosis ad Kabousie alt. 3500 pedes, Murray, 54!. (Sent

by Mr. MacOwan to Herb. Kew.)’. It is remarkable to

note that Baker (1874) described this species as hav-

ing "folia . . . glabra ’ and "stylus filiformis . . . stigmate

capitato obscure trilobato\ the latter character being

the basis to create his new monotypic section, Albuca

sect. Leptostyda. However, the original material present

at K — Colesberg, Shaw s.n. (K0002573 12); Vaal rivier,

Shcn\’ s.n. (K000257313); in graminosis ad Kabousie,

Murray 54 (K000257314) — include plants that match

the traditional concept of^. shawii, and they indeed bear

long stipitate glands in most of their parts that are visible

to the naked eye. Moreover, Murray 54 (K000257314)

has a typically obpyramidal style, whereas other gyn-

oecia are thinner (resembling a narrow style), probably

due to desiccation. This is congruent with comments

by Hilliard & Burtt (1982: 282) on this same subject.

Gynoecium differences with regard to A. tenuifolia are

evident when living plants are compared (Figure 2).

All populations of A. shawii we observed in the wild

showed constantly obpyramidal styles. Furthermore, two

of the synonyms of A. shawii that are currently accepted

(e.g. A. trichophylla Baker and A. minima Baker) were

described as having pubescent, not glandulose leaves,

whilst their types show plants with long stipitate glands.

These incongruences in the protologues are similar to

that concerning A. tenuifolia, as cited before.

According to the data presented above both taxa

should be regarded as distinct at the species rank, as

shown below.

TAXONOMY

Albuca tenuifolia Baker, Refugium Botanicum

[Saunders]: t. 335 (1872). Lectotype, designated by

FIGURE 2. — Flower dissections of A. tenuifolia Baker (A, Eastern Cape, Somerset East, summit of Boschberg, Clark & Mariinez-Azorin 23 GRA)

and Albuca shawii Baker (B, Eastern Cape, Linedrift, near Keiskamma River Bridge on N2, Martinez-Azorin, Dold & Martinez-Soler 520

GRA). Outer tepals and stamens above; inner tepals and stamens below; lateral views of gynoecia. Scale bar: 1 cm.

196

Bothalia 42,2 (2012)

Martinez-Azorin et al.: 467 (2011): Baker: t. 335 (1872).

Epitype, designated by Martinez-Azorin et al.: 467

(2011): South Africa, Eastern Cape, Somerset East, sum-

mit of Boschberg, northeast of Bloukop, in shallow soil

on rocky places, 1 574 m, 10 May 2010, V.R. Clark &

M. Martinez-Azorin (GRA, epi.; ABH, K, PRE, isoepi.)

Diagnostic characters: Albuca tenuifolia can be eas-

ily identified by its hypogeal and proliferous, irregularly

compressed bulbs; filiform and straight leaves that bear

minute and sessile papillae (only visible under micro-

scope); glabrous peduncle and pedicels; erect pale yel-

low-green flowers; strongly hooded inner tepal tips; all

stamens bearing fertile anthers; subglobose ovary with

divergent paraseptal crest; and a narrowly obpyramidal

trigonous style.

The characteristic form of the inner tepal tips,

strongly hooded and resembling a mitre (Figure 2a)

together with the erect flowers, place A. tenuifolia in A.

subg. Mitrotepalum (= A. sect. Branciona). For further

details and discussion on subgeneric arrangement of

Albuca see Muller-Doblies (1987, 1995).

Albuca shawii Baker in Journal of Botany (London)

12: 367 (1874). Lectotype, designated by Hilliard &

Burtt: 281 (1982): South Africa, Northern Cape, Coles-

berg, Shaw s.n. (K000257312, lecto. — photo!).

Diagnostic characters: Albuca shawii is characterized

by its hypogeal, ovoid, and mostly solitary bulbs; fili-

form leaves with usually coiled apexes; leaves, pendun-

cle, and pedicles with evident, stipitate glands; nodding

yellow-green flowers; inner tepals with a slight apical

hood and a triangular structure facing downwards; only

the three inner stamens bearing fertile anthers, the out-

ers commonly lacking anthers; oblong ovary with almost

straight paraseptal crests; and the shorter and widely

obpyramidal trigonous style.

The inner tepals tips of A. shawii are neither strongly

hooded nor hinged (Figure 2b), a character which,

together with the nodding flowers, place this species in

A. subg. Falconera (= A. sect. Falconera) (cf Miiller-

Doblies 1987, 1995) The species is the type of A. sect.

Trianthera U.Mtill.-Doblies (= A. ser. Trianthera

(U.Miill.-Doblies) J.C. Manning & Goldblatt), character-

ized by the presence of only three fertile anthers.

ACKNOWLEDGEMENTS

This work was partly supported by the Fundacion

Ramon Areces (Spain). We thank the curators of the her-

baria ABH, BOL, BNRH, GRA, J, K, KEI, KMG, NBG,

NH, NU, PEU, PUC, UFH, and WfND, for allowing us

to study their Albuca collections. Thanks to P. Wilkin,

O. Weber and L. Walsingham (Royal Botanic Gardens,

Kew) for supplying images of some Albuca collections

at Kew. We also thank F. Martinez-Flores who kindly

helped us with the figures of Albuca leaves.

REFERENCES

BAKER, J.G. 1872. Albuca tenuifolia. In W.W. Saunders, Refugium

botanicum 5(2) t. 335. Van Voorst, Paternoster Row, London.

BAKER, J.G. 1873. Revision of the genera and species of Scilleae and

Chlorogaleae. Journal of the Linnean Society. Botany 13(68):

209-292.

BAKER, J.G. 1874. Description of new species of Scilleae and other

Liliaceae. Journal of Botany. British and Foreign 12: 363-369.

BAKER, J.G. 1897. Albuca. In W.T. Thiselton-Dyer, Flora Capensis 6:

451-462. Reeve & co., London.

HILLIARD, O.M. & BURTT, B.L. 1982. Notes on some plants of

southern Africa, chiefly from Natal: IX. Notes front the Royal

Botanic Garden, Edinburgh 40: 247-298.

MCNEILL, J., BARRIE, F.R., BURDET, H.M., DEMOULIN, V,

HAWSWORTH, D.L., MARHOLD, K., NICHOLSON, D.H.,

PRADO, J., SILVA, PC., SKOG, J.E., WIERSEMA, J.H. &

TURLAND, N.J. (EDS.). 2006. INTERNATIONAL CODE OF

BOTANICAL NOMENCLATURE (VIENNA CODE) ADOPT-

ED BY THE SEVENTEENTH INTERNATIONAL BOTANICAL

CONGRESS VIENNA. AUSTRIA. JULY 2005. GARTNER VER-

LAG, RUGGEL, LIECHTENSTEIN [REGNUM VEGETABILE

146].

MANNING, J.C. & GOLDBLATT, P. 2003. Albuca. In G. Germishui-

zen & N.L. Meyer, Plants of Southern Africa: an annotated

checklist. Strelitzia 14: 1054-1056. National Botanical Institute,

Pretoria.

MARTINEZ-AZORIN, M., CLARK, V.R., CRESPO, M.B., DOLD

A.P. & BARKER N.P. 2011. The rediscovery oi Albuca tenuifo-

lia, an orophilous species from the eastern Great Escarpment in

South Africa. Nordic Journal of Botany 29: 465-470.

MULLER-DOBLIES, U. 1987. Infrageneric taxonomy of y4/6i/ca (Hya-

cinthaceae). Abstract XIV International Botanical Congress p.

332. Berlin.

MOLLER-DOBLIES, U. 1995. Enumeratio Albucarum (Hyacinthace-

ae) Austro-Africanarum adhuc cognitarum. 2. Subgenus Falcon-

era (Salisb.) Baker emend. U.M.-D 1987. Feddes Repertorium

106: 353-370.

MULLER-DOBLIES, U. 2012. The enigma of Albuca tenuifolia Baker

resolved. Feddes Repertorium 121: 285-298.

THIERS, B. 2012. Index herbariorum: a global directory of public her-

baria and associated staff, [continuously updated]. New York.

Bot. Gard. Virt. Herb, <http://sweetgum.nybg.org/ih/>, accessed

January 2012.

Mario Martinez-Azorin*, Manuel B. Crespo* & Anthony P. Dold**

* CIBIO (Instituto Universitario de la Biodiversidad), Universidad de

Alicante, PO. Box 99. E-03080 Alicante, Spain, mmartinez@ua.es (au-

thor for correspondence).

" Selmar Schonland Herbarium, Department of Botany, Rhodes Uni-

versity, Grahamstown 6140, South Africa.

MS. received: 2012-01-31.

ITEACEAE

TAXONOMIC AND NOMENCLATURAL NOTES ON THE GENUS CHORISTYLIS

Since the description of the sub-Saharan African

Choristylis and its only species, C. rhamnoides by Har-

vey in 1842, the family position of this woody shrub or

small tree has been uncertain. We review the taxonomic

status and family position of the genus and summa-

rize its nomenclatural history. Harvey (1862), Hooker

(1865), Mildbraed (1934), Boutique (1964) and Liben

(1969) referred Choristylis to Saxifragaceae, and Engler

Bothalia 42,2 (2012)

197

(1891, 1930) placed it in Saxifragaceae subfamily Escal-

lonioideae. Other authors, including Hutchinson (1969),

Verdcourt (1973, 1983), Brummitt (1992), Jordaan

(2000), Letter (2002), Coates Palgrave (2002) and Boon

(2010) included Choristylis in the Escalloniaceae. Airy

Shaw (1973) was the first to associate Choristylis with

Itea in the small family Iteaceae J.G.Agardh (sweet-

spire family), a step followed by Takhtajan (1996) and

Kubitzki (2006). This is the family placement followed

here.

Saxifragaceae are exclusively herbaceous with exstip-

ulate leaves. Escalloniaceae are trees with exstipulate

leaves and a parietal placentation, whereas in the woody

Iteaceae the leaves are always stipulate, placentation is

axile and the pollen grains are unusual in being 2-porate

and subisopolar (Erdtman 1955; Agababian 1960). Pol-

len in Escalloniaceae is shed as monads and the grains

are isopolar and 3-5-colporate. Iteaceae also differ from

Escalloniaceae in their bitegmic and crassinucellate

ovules. Escalloniaceae have unitegmic and tenuinucel-

late ovules. The crassinucellate condition was deter-

mined in both Itea and Choristylis (Mauritzon 1933,yz<7e

Kubitzki 2006). APG III (2009) place Escalloniaceae

in an order of its own, the Escalloniales. Fossil flowers

resembling those of Iteaceae and dating from the Turo-

nian stage of the Late Cretaceous have been found in the

Raritan Formation, New Jersey, and leaves ascribed to

Iteaceae and dating to the Eocene have been described

from the Klondike Mountain Formation, Washington

(Hermsen et al. 2003; Friis et al. 2011).

Molecular analyses place Itea and Choristylis in the

core Saxifragales, sister to Pterostemon Schauer (Pter-

ostemaceae) (Savolainen et al. 2000; Fishbein et al.

2001). Pterostemonaceae (Engl.) Small is a monogeneric

family of two species of shrubs native to tropical and

subtropical Mexico. This family, formerly associated

with Grossulariaceae s.l., Escalloniaceae, and Hydrange-

aceae by some authors, is currently included in Iteaceae

by APG III (2009). Studies done by Bohm et al. (1999)

showed that the flavonoid profiles of Pterostemon con-

sist of C-glucosylflavones and 3-O-glycosides of querce-

tin, which resembles very closely that observed in Itea.

The presence of C-glucosylflavones in these two genera

provides additional support for the closeness of their

relationship. In contrast, Saxifragaceae, Crassulaceae,

and other members of the Saxifragales are characterized

by flavonol production (Bohm et al. 1988).

Kubitzki (2006) relegates Choristylis to the syn-

onymy of Itea, a genus with about 1 9 species, of which

15 species (ten endemic) occur in south and southeast

Asia and Malaysia (Jin & Ohba 2001), one in Paki-

stan (Siddiqi 2005), one in Japan (Ohwi 1965), one in

North America (Morin 2009) and one species in Africa

(Verdcourt 1973, 1983). Members of Iteaceae have spi-

rally-arranged or alternate, glandular-dentate, serrate to

spinose leaves with superposed axillary buds, paniculate

to racemose inflorescences with rather small 5-merous

flowers, and septicidally dehiscent fruits, with the valves

often remaining attached by the stigma. The flowers

have a hypanthium, the corolla is valvate, and there are

only two carpels. Flowers of Saxifragaceae have four or

five free sepals and petals, with stamens as many as or

twice as many as the sepals, and the pistil consisting of

2-5 united carpels. Although the carpels of Iteaceae are

free initially, they become partly or completely united

along the style (Ge et al. 2002; Kubitzski 2006).

The African Choristylis is notably disjunct from other

members of Itea, which are exclusively Northern Hemi-

sphere in distribution, extending from south and south-

east Asia from the Himalayas to China, Taiwan, Japan,

Java, the Philippines, and southeast United States of

America. Choristylis, however, is similar in habit, leaf,

flower, fruit, and seed characters to Itea. Members of

Itea have alternate or spirally arranged, glandular ser-

rate or dentate to spinose-dentate leaf margins with the

lateral or tertiary veins ending at the margin in a gland

or spine (the same condition as in Choristylis) to rarely

entire. The leaf venation is seemingly 3-veined from

or near the base in Choristylis and pinnately veined in

all species of Itea. Both genera have similar, small, 5-

merous, creamy white or greenish yellow flowers

borne in axillary or terminal panicles or racemes, often

superposed in groups of two or three. In Choristylis

the inflorescences are shorter than the leaves, up to 45

mm long, whereas most Itea species have long, droop-

ing, elongated panicles or racemes, up to 60-200 mm

long, much longer than the leaves (Jin & Ohba 2001;

Siddiqi 2005; Morin 2009). The flowers in both genera

have a turbinate or obconic calyx-tube adnate to the base

of the ovary and with five lobes; the petals are valvate

and persistent; the stamens are inserted at the margin

of a hypogynous disc and are included or exserted; the

ovary is 2-locular, superior to half-inferior; the styles

are undivided or ± deeply divided, but apically coherent

at anthesis. The fruit is a septicidally dehiscent capsule

with a persistent perianth in both genera. Given the gen-

eral morphological similarity between the two genera

and their identical pollen we agree with Kubitzki (2006)

that the range of variation of characters, such as ovary

position (nearly superior to half inferior), the degree of

fusion of the styles (undivided or ± deeply divided,) and

ramification and length of inflorescences (terminal or

axillary panicles or racemes), makes the maintenance of

Choristylis impossible.

Bentham (1865) kept Choristylis and Itea separate

when only a few species of Itea were known, and stated

that the one Itea species from America (/. virginica L.)

and one from Japan (I. japonica Oliv.) are deciduous and

that their seeds are oblong and flattened, distinguishing

them from the Asian species, which are evergreen trees

and their seeds are fusiform. Since then many more

species of Itea have been described and Engler (1891,

1930) consequently took up these characters mentioned

by Bentham and used them to distinguish two sections

in Itea, namely sect. Deciduae Engl, for the American

and Japan species, and sect. Sempervirentes Engl, for the

Asian species.

Although Kubitzki (2006) argued that Engler ’s sec-

tions (1891, 1930) in Itea could not be upheld with the

inclusion of Choristylis within Itea, the seed morphol-

ogy of Choristylis differs enough from the other species

to warrant a section of its own in the genus Itea. Chori-

stylis is an evergreen scrambling shrub or tree and the

seeds are irregularly obovoid or oblong-obovoid, flat-

tened, sculptured and slightly curved (Verdcourt 1973,

tab. 1,13; Kubitzki 2006, fig. 70G). We accordingly

Bothalia 42,2 (2012)

describe a new section within Itea to accommodate the

single African species.

TAXONOMY

Itea L. sect. Choristylis (Harv.) Jordaan, sect. nov.

et stat. nov. Choristy’lis Harv.: 19 (1842). Similar to sect.

Sempef^virentes Engl, in its evergreen habit but differ-

ing in its leaves seemingly three-veined from or near

the base, short inflorescences less than 45 mm long, and

seeds which are irregularly obovoid or oblong-obovoid,

flattened, sculptured and slightly curved.

Type species: Itea rhamnoides (Harv.) Kubitzki.

{=Choristylis rhamnoides Harv.)

Itea rhamnoides (Harv.) Kubitzki in K. Kubitzki,

The families and genera of vascular plants 9: 203

FIGURE I . — Jlea rhamnoides. A, flowering twig, x I; B, leaf with irregular glandular-serrated margin, x 2; C, young flower, x 15; D, mature flower,

X 15; E, fruit, x 15. A & C taken from Thode A3 1 8 {PREy, B taken from Dahlsirand 2669 {PRE); D taken from Jordaan 3938 (PRE); E taken

from Theron 3648 (PRU), Artist: Daleen Roodt.

Bothalia 42,2 (2012)

199

(2006). Choristylis rhamnoides Harv.: 19 (1842), 308

(1862). Type: South Africa, Eastern Cape, Katberg,

Brownlee s.n. (TCD, holo.-Aluka image!; K, iso.).

Choristylis shirensis Baker f: 13, t. 3/1-6 (1894).

Type; Malawi, Mt. Milanji [Mulanje], 1891, Whyte S3

(BM-Aluka image!, lecto., designated here; Z-Ztirich

Virtual Herbarium!, isolecto.). [Whyte 53 in the British

Museum is chosen as the lectotype as the best represent-

ative specimen among the available syntypes].

C. iilngiirensis Mildbr.: 191 (1934). Type: Tanganyika

[Tanzania], Uluguru Mts., southern slope of Lukwangule

Peak, August 1933, Schliehen 4293 (B B100165111-

Aluka image!, lecto., here designated). [The type col-

lection of Schliehen 4239 on which Mildbraed (1934)

based his name consists of two specimens, of which

number B100I65111 is the better specimen with more

flowering material and therefore chosen as lectotype].

Evergreen scrambling shrub or small tree up to 7

m tall, with slender trailing branches, single- or multi-

stemmed; stems slender, diverging from base, angu-

lar-ribbed; young branchlets often purplish, covered

with short hairs. Bark on stems splitting longitudinally.

Leaves spirally arranged, glossy green above, paler and

dull below, hairless or with sparse hairs on the main

veins below and often with domatia; lamina ovate or

oblong to ovate-lanceolate, 15-103 x 9-55 mm, apex

acute to shortly acuminate or rarely rounded, base

cuneate to unequally rounded; margins sharply and

closely irregular glandular-serrate; seemingly 3-veined

from base, but otherwise pinnately veined; midrib and

main veins sunk above, prominent below, fine tertiary

reticulate venation conspicuous; petioles often purple,

up to 20 mm long, pubescent. Stipules small, decidu-

ous. Inflorescences many-flowered panicles, shorter than

leaves, finely hairy, up to 45 mm long, in dense clusters;

bracts and bracteoles narrowly triangular, 0.5-1. 5 mm

long; peduncles 0-12 mm long. Flowers very small, reg-

FIGURE 2. — Distribution of Itea rhamnoides, based on specimens

housed in the National Herbarium, Pretoria, and in literature

(Liben 1969 & Verdcourt 1973, 1983).

ular, hermaphrodite or polygamous, 5-merous, creamy

white or greenish yellow, sweetly scented; pedicels

1- 2(-5) mm long. Calyx tube (hypanthium) short, ± 1.5

X 1.5 mm, adnate to ovary; lobes triangular, 0. 5-2.0 x

0.3-0.5(-l .0) mm at base, ciliate. Petals ovate, 1.5-3. 5

X 1-2 mm at base, 3-veined, hairy. Stamens slightly

exserted, hairy; anthers ovoid, hairy. Disc annular.

Ovary’ half-inferior, bicarpellate, carpels free initially,

2- locular; ovules numerous, biseriate, with axile placen-

tation; styles 2, subulate, adherent or diverging, 0. 5-1.0

mm long, yellowish, turning brown; stigmas ± 0.5 mm

diam., free or sometimes cohering for a time. Print a

septicidally dehiscent capsule, campanulate to turbinate

below, conical above, 3-5(-6) x 3 mm, brownish, finely

hairy, slightly ribbed, leathery. Seeds obovoid or oblong-

obovoid, flattened, ± 1.0 x 0.5 mm, reticulate, brownish.

Figure 1.

Diagnostic characters'. Itea rhamnoides can be rec-

ognized by its leaves, which are spirally arranged, ±

3- veined at or near the base, with glandular toothed mar-

gins and conspicuous tertiary reticulate venation. The

ovary is half-inferior with two diverging styles, whence

is derived the name Choristylis, from the Greek word

choris = separate and stylis = style. The valves of the

septicidally dehiscent capsules often remain attached by

the stigma.

Distribution and habitat'. Itea rhamnoides grows in

the eastern parts of southern Africa, from the Katberg,

Hogsback, Keiskammahoek, and Kentani in the Eastern

Cape, northwards through KwaZulu-Natal, Swaziland

and Mpumalanga to the Soutpansberg in the north. From

here it extends further northwards through Zimbabwe,

Mozambique, and Malawi along the mountains to the

Democratic Republic of the Congo, Burundi, Uganda,

and Tanzania along the East African Rift, but seems to

be absent from the more recent volcanic mountains in

East Africa (Kenya and Ethiopia) (Verdcourt 1983) (Fig-

ure 2). The distribution coincides with that of the archi-

pelago-like Afromontane Region of Endemism (White

1978, 1983). It grows in forest, forest margins, riverine

and valley forests, open hillsides, often along streams at

medium to high altitudes.

Synonymy. Harvey (1862) considered Baeobot-

tys rufescens E.Mey. to apply to Choristyles rham-

noides, which De Candolle (1844) placed in the genus

Maesa, based on a Drege specimen, and Maesa palus-

tris Hochst. (1844), based on a Krauss specimen (Krauss

152), under Choristylis rhamnoides. These synonyms,

however, belong to Maesa lanceolata Forssk. (Dyer

1963). Verdcourt (1973) was the first to place the tropi-

cal African Choristylis shirensis Baker f. (1894) from

Malawi and C. ulugurensis Mildbr. (1934) from Tanza-

nia in synonymy under C. rhamnoides. The name Chori-

stylis virescens Baker f, which appears in The Interna-

tional Plant Names Index [IPNI] as published on the

same page where Baker described Choristylis shirensis

(Baker 1894) seems to be fictitious (Verdcourt 1983).

We have checked Baker (1894) and also cannot find any

mention of C. virescens. We conclude that it is a spelling

error for C. shirensis.

200

Bothalia 42,2 (2012)

ACKNOWLEDGEMENTS

I would like to thank Hester Steyn for preparing the

distribution map, and the artist Daleen Roodt for the line

drawing.

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VERDCOURT, B. 1973. Escalloniaceae. In R.M. Polhiil, Flora of tropi-

cal East Africa: 1-3. Crown Agents for Oversea Governments

and Administrations, London.

VERDCOURT, B. 1983. Escalloniaceae. In E. Launert, Flora zambe-

siaca 7,1: 1-3. Flora Zambesiaca Managing Committee, Edin-

burgh.

WHITE, F. 1978. The Afromontane Region. In M.J.A. Werger, Bioge-

ography and ecology) of southern Africa. Junk, The Hague.

WHITE, F. 1983. The vegetation of Africa: a descriptive memoir to

accompany the UNESCO, AETFAT, UNSO vegetation map of

Africa, Paris.

Websites accessed

ALUKA LIBRARY. http.7/plants,jstor.org/ [accessed September 2010].

ANGIOSPERM PHYLOGENY WEBSITE (APweb). www.mobot.org/

MOBOT/Research/APweb/ [accessed August 2009].

EFLORAS. http://www.efloras.org/ [accessed March 2012].

HERBARIA OF THE UNIVERSITY AND ETH ZURICH (Z, ZT &

ZSS). http://www.zuerich-herbarien.unizh.ch/ [accessed Febru-

ai7 20ll].

INTERNATIONAL PLANT NAMES INDEX [IPNI]. Published on the

Internet http://www.ipni.org/ [accessed July 2009].

TROPICOS. http://www.tropicos.org/ [accessed February 2011].

VIRTUAL HERBARIUM OF THE NATURAL HISTORY MUSEUM

VIENNA (W) AND INSTITUTE OF BOTANY, UNIVER-

SITY OF VIENNA (WU) http://herbarium.botanik.univie.ac.at/

[accessed February 2011].

M. JORDAAN'

' National Herbarium, South African National Biodiversity Insti-

tute, Private Bag XIOI, 0001, Pretoria. / Student affiliation: Depart-

ment of Plant Science, University of Pretoria, 0002, Pretoria. E-mail:

m.jordaan@sanbi.org. za.

MS, received: 2012-01-10.

Bothalia 42,2 (2012)

201

NYCTAGINACEAE

TAXONOMIC STATUS OF COMMICARPUS FRUTICOSUS

Commicarpus squarrosus (Heimerl) Standi, is

endemic to the Kunene, Erongo, Hardap, and Karas

regions of Namibia (Struwig 2012). A closely related

taxon, C. fruticosus Pohnert, is endemic to the Tsaris and

Naukluft Mountains (north of the town of Zaris in the

Hardap region of Namibia). This name was subsequently

treated by Schreiber (1969) as a synonym of C. squarro-

sus in the Prodromus einer Flora von Sudwestafrika for

unspecified reasons. The view of Schreiber is not widely

supported and C. fruticosus is generally recognized as

a separate species (Nowicke 1970; Craven 1999; Ger-

mishuizen & Meyer 2003; Klopper el al. 2006; African

Plant Database 2012).

No workable diagnostic morphological differences

have hitherto been documented between C. fruticosus

and C. squarrosus, and even the original descriptions

of the two species (Heimerl 1896; Pohnert 1953) are

noticeably similar. The exception being that the original

description of C. fruticosus, Pohnert (1953) suggests that

the species produces false whorls in its inflorescence.

This was refuted as a diagnostic character (Struwig

2012), as this phenomenon is often present throughout

the range of C. squarrosus. Although the flowers of C.

fruticosus were described as white, greyish-white to pur-

ple, our collections of C. fruticosus from the Naukluft

Mountains all had dark purple flowers. Flower colour

in the widespread C. squarrosus varies from light pur-

ple to pink. This suggests that the flower colour of the

Naukluft Commicarpus species is variable and overlaps

with that of Commicarpus squarrosus . Flower colour in

Nyctaginaceae is generally a weak taxonomic character

and not considered useful in distinguishing between spe-

cies (Struwig 2012).

Anthocarp morphology, on the other hand, is consid-

ered to be the most taxonomically useful character in

distinguishing genera and species of the Nyctaginaceae

(Bohlin 1988; Douglas & Manos 2007) and has been

proven effective for the southern African Commicarpus

species (Struwig et al. 2011).

Anthocarp morphology in Commicarpus squarrosus

and C. fruticosus is identical (Struwig 2012); fusiform

and ten-ribbed, (4-)5(-6) x (l-)1.8(-2) mm, glabrous,

with the apex surrounded by five stalked glands alter-

- '4 ^

- .0

. ■ Vj, . , ' I’A

O I®

FIGURE 1. — Anthocarps of A, Commicarpus fruticosus (Struwig 164)\

and B, Commicarpus squarrosus (Struwig 36). Scale bars 1 mm.

Material was collected and preserved in 4% paraformaldehyde

and micrographs were taken with a Nikon Digital Camera DXM

1200 F fitted on a Nikon SMZ 1500 stereomicroscope.

nating with five sessile glands and the surface below the

apex characterized by scattered sessile glands (Figure 1 ).

Commicarpus fruticosus should therefore be re-instated

as a synonym of C. squarrosus on the basis of similari-

ties in inflorescence structure and anthocarp morphol-

ogy. Table 1 summarises the presence of the genus Com-

micarpus in southern Africa.

Commicarpus squarrosus (Heimerl) Standi., in

Contributions from the United States National Herbar-

ium 18: 101 (1916). Boerhaavia squarrosa Heimerl: 813

(1896). Type: Namibia, ‘Gross-Namaland, Rehoboth’,

Fleck 241 A (Z, holo. — digital image!).

Commicarpus fruticosus Pohnert: 337 (1953). Type:

Namibia, ‘Tsarisberge, Zebraschlucht’, Walter& Walter

2095 (M, holo.-Aluka image!, website accessed 25 Aug.

2011; WIND, iso.!).

Additional specimens

NAMIBIA. — 1913 (Sesfontein): Khowarib Rest Camp, hills sur-

rounding rest camp, (-BD), 9 Feb. 2009, Struwig 45 (PUC, WIND).

2014 (Khorixas): Twyfelfontein Lodge, hillslope behind lodge, SW of

TABLE 1. — Commicarpus species occurring in the Flora of southern Africa countries. E, endemic to southern Africa (Struwig 2012).

202

Bothalia 42,2 (2012)

main building, high up on slope amongst big boulders, (-CB), 7 Feb.

2009, Stniwig 41 (PUC, WIND). 2114 (Uis): Numas Gorge, high up

in kloof, underneath Acacia montis-usti behind big boulders, (-BA),

6 Feb. 2009, Striiwig 39 (PUC, WIND); Brandberg Mountain, on

top, Baswald Rinne Area, (-BC): 19 May 1977, Craven 490 (PUC,

WIND). 2115 (Karibib): Klein Spitzkoppe, against mountain slope

amongst rocks, (-CC), 5 Feb. 2009, Struwig 36 (PUC, WIND). 2214

(Swakopmund): Swakopmund district, 63 miles SE of Walvis Bay in

Namib area, (-DA), 2 Mar. 1965, Barnard 85 (PRE, WIND), 2215

(Trekkopje): Tsaobis Leopard Farm, S of Karibib, (-DD), 21 Feb.

1990. Hardy 7017 (PRE, WIND). 2416 (Maltahdhe): Naukluft Moun-

tains at Billlspoort, (-AA), 16 Dec. 1947, Rodin 2833 & Strey 2132

(BOL); Cl 4, Naukluft Mountains, mountain slope behind the river, (-

AA), 13 Feb. 2009, 7 Apr. 2010, 8 Apr. 2010, Struwig 59. 160. 163

(PUC, PRE, WIND); Farm Tsais-Maltahohe, (-AB), 16 May 1978,

Muller & Tilson 894 (PRE, WIND); C19. Tsaris Mountains, (-AB), 8

Apr. 2010, Struwig 164 (PUC, WIND); Maltahdhe, Earm Mooirivier

MAE 160, on S-facing slopes, (-CA), 11 Apr. 1980, Muller 1362

(PRE, WIND). 2616 (Aus): Kuibis, (-DD), 1 Mar. 1912, Range 1283

(BOL).

ACKNOWLEDGEMENTS

The curators of the following herbaria are acknowl-

edged for providing access to study material: BOL,

PRE, PUC, and WIND (acronyms according to Holm-

gren el al. 1990). The North-West University and

National Research Foundation provided financial sup-

port.

REFERENCES

AFRICAN PLANT DATABASE. 2012. Version 3.3. Conservatoire

et Jardin botaniques de la Ville de Geneve and South African

National Biodiversity Institute, Pretoria, http://www.ville-ge.ch/

musinfo/bd/cjb/africa/ [accessed Apr. 2012].

BOHLIN, J.-E. 1988. A monograph of the genus Colignonia (Nyctagi-

naceae). Nordic Journal of Botany 8: 231-252.

CRAVEN, P (ed.). 1999. A checklist of Namibian plant species. South-

ern African Botanical Diversity Network Report No. 7. SAB-

ONET, Windhoek.

DOUGLAS, N.A. & MANOS, P.S. 2007. Molecular phytogeny of

Nyctaginaceae: Taxonomy, biogeography and characters asso-

ciated with a radiation of xerophytic genera in North America.

American Journal of Botany 96: 856-872.

GERMISHUIZEN, G. & MEYER, N.L (eds.). 2003. Plants of southern

Africa: an annotated checklist. Strelitzia 14. National Botanical

Institute, Pretoria.

HEIMERL, A. 1 896. Nyctaginaceae. In E. Autran, Bulletin de I 'Herbier

Boissier 1: 813. Imprimerie Romet, Geneve.

HOLMGREN, P.K., HOLMGREN, N.H. & BARNETT, L.C. 1990.

Index Herbariorum. part 1: the herbaria of the world. 8"' edn.

New York Botanical Garden, New York.

KLOPPER, R.R., CHATELAIN, C., BANNINGER, V., HABASHI, C.,

STEYN, H.M., DE WET, B.C., ARNOLD, T.H., GAUTIER, L.,

SMITH, G.F. & SPICHIGER, R. 2006. Checklist of the flower-

ing plants of sub-Saharan Africa. An index of accepted names

and synonyms. South African Botanical Diversity Network

Report No. 42. SABONET, Pretoria.

NOWICKE, J.W. 1970. Pollen morphology in the Nyctaginaceae.

Grana 10: 79-88.

POHNERT, H. 1953. Commicarpus fallacissimus (Heimerl) Pohnert. In

K. Suessenguth, Mitteilugen der Botanischen Staatssammlung

Miinchen. 8:337. Munchen.

SCHREIBER, A. 1969. Nyctaginaceae. In H. Merxmiiller, Prodromus

einer Flora von Siidwestafrika 25. J. Cramer, Lehre.

STANDLEY, P.C. 1916. Systematic investigations of tropical American

plants. Contributions from the United States National Herbari-

um 18: 101. Government Printing Office, Washington.

STRUWIG, M., SIEBERT, S.J. & KLAASSEN, E. 201 1 . Nyctaginace-

ae. Notes on Commicarpus Standi, in southern Africa, including

a new record for Namibia. Bothalia 4\: 289-292.

STRUWIG, M. 2012. A systematic study q/ Boerhavia L. and Commi-

carpus Standi. (Nyctaginaceae) in southern Africa. Ph.D. thesis,

North-West University, Potchefstroom.

M. STRUWIG'" and S.J. SIEBERT*

‘A.P. Goossens Herbarium, School of Environmental Sciences and De-

velopment, North-West University, Private Bag X6001, 2520 Potchef-

stroom, South Africa.

" Author for correspondence: madeleen.struwig@nwu.ac.za.

MS. received: 2012-04-24.

POACEAE

THE TAXONOMIC AND CONSERVATION STATUS OF AGROSTIS ERIANTHA VAR. PLANIFOLIA

INTRODUCTION

Agrostis ericmtha Hack. (1904) is a tufted, rhizoma-

tous perennial that grows in wetlands of Swaziland,

Lesotho, Limpopo, Mpumalanga, Gauteng, Free State,

KwaZulu-Natal, and Eastern Cape. This relatively

rare grass appears to be sensitive to disturbance and is

mainly found in pristine habitats. In 1945, Goossen

Papemdorf described a form of the species collected by

Pole-Evans on the farm Doornkloof, Irene, as A. erian-

tha Hack.var. plcmifoHa Gooss. & Papemdorf. The main

diagnostic character used to distinguish the two varieties

was the length of the callus hairs as shown in Figures 1

and 2. In Agrostis ericmtha var. ericmtha, the callus hairs

are up to one third the lemma length while in var. plaai-

folia, the callus hairs are up to half the lemma length.

Another suggested difference was in the leaf blades,

which are said to be folded in var. eriantha and flat in

var. plattifolia. Other possible differences are discussed

in the results section.

The only known collections of A. eriantha var. plani-

folia are the type specimen and another Pole-Evans

collection from Irene, eollected two days after the type

specimen. However, examination of Pole-Evans’ regis-

ter shows that he made mistakes with localities and was

also not consistent with dates, casting doubt on whether

the second specimen was really another collection rather

than a duplicate of the type. Unfortunately, as with the

Type specimen, there is very little information. Since

this plant was first discovered, forms with callus hair

length equal to that of the type specimen have never

been found again, despite repeated searches at the type

locality and nearby habitat.

Various unanswered questions concerning A. eriantha

var. planifolia have made it difficult for an assessment

of the eonservation status of this plant to be made. Since

it has only been collected once, it remained uncertain

whether it is extremely limited in distribution and still

awaiting re-discovery, or alternatively, extinct. However,

Bothalia 42,2 (2012)

203

FIGURE 1. — Agrostis eriantha subsp. planifolia, Pole-Evans 666 (PRE). A, spikelet; B, lemma. Scale bar: A, B, 200 pm. Photographer: Caroline

Mashau.

FIGURE 2. — Agrostis eriantha subsp. eriantha, Smook 5026 (PRE). A, spikelet; B, lemma. Scale bar: A, B, 200 pm. Photographer: Caroline

Mashau.

it is also possible that this taxon is merely an aberrant

form of A. eriantha. In recent years, development pro-

posals in the vicinity of this taxon have been compli-

cated by its potential (but unconfirmed) presence, a situ-

ation which has financial implications for developers.

Thus it is of great importance to clarify the taxonomic

and conservation status of this taxon.

The current conservation status of this variety (Rai-

mondo et al. 2009) is Data Deficient (DD). Additionally

it is flagged ‘T’, indicating taxonomic uncertainty (Vic-

tor 2006).

METHODS

Fieldwork was conducted at the type locality and sur-

rounding areas in Gauteng. The type locality has been

transformed and no Agrostis eriantha remains there. The

two closest wetlands to the type locality that are in a rel-

atively reasonable condition are Rietvlei Nature Reserve

and the Grootfontein Agricultural Holdings. Specimens

of A. eriantha were collected from both of these sites.

Callus hairs of the specimens were compared with those

of A. eriantha var. planifolia. In addition, the callus

hairs of 94 specimens of Agrostis eriantha var. eriantha

housed in the National Herbarium (PRE) were investi-

gated to determine variation in callus hair length within

this taxon.

As a comparison, callus hair variation in Agrostis

lachnantha Nees, a closely related and sympatric spe-

cies, was investigated to determine the consistency of

this character. Samples from 11 specimens were inves-

tigated.

RESULTS

The comparison shows that the morphological char-

acteristics of specimens collected near the type locality

match the type and description of A. eriantha var. eri-

antha. No specimen of A. eriantha var. eriantha has cal-

lus hairs quite as long as A. eriantha var. planifolia, but

some variation in length was found (Table 1).

Variability of callus hair length between different

specimens of Agrostis lachnantha was investigated to

assess the reliability of callus hair length as a character.

This investigation revealed that callus hair length varied

up to one third the lemma length not only between spec-

imens but also within the same specimen.

204

Bothalia 42,2 (2012)

TABLE 1 . — Differences between Agrostis erianiha var. eriantha and A. eriantha var. planifolia

DISCUSSION AND CONCLUSION

Agrostis lachnantha was divided into two varieties, A.

lachnantha var. lachnantha and A. lachnantha var. gla-

bra on the basis of hairiness of the lemma by Goossen

& Papendorf (1945). However it was later found, and

confirmed in this investigation, that hairs on the lemma

are variable in length and cannot reliably be used to dis-

tinguish varieties. The variety was therefore reduced to

synonymy under A. lachnantha (Gibbs Russell et al.

1990).

The results of our investigations suggest that vari-

ation in callus hair length is not a reliable character to

use to distinguish between taxa in Agrostis eriantha.

Given that A. eriantha var. planifolia has never been

recollected (with the exception of one other specimen

from the type locality), it is probable that it is an aber-

rant form. Agrostis eriantha var. planifolia is therefore

reduced to synonymy under A. eriantha. This species is

a widespread grass and the conservation status is con-

firmed to be Least Concern.

Agrostis eriantha Hack, in Vierteljahresschrift der

Naturforschenden Gesellschaft in Zurich 49: 172 (1904).

Syntypes: South Africa, [Gauteng], ‘in humidis prope

Pretoria’, Jan. 1894, Schlechter 4144 (PRE, syn.!);

[Eastern Cape], ‘in collibus prope Middleburg’, Dec.

1893, Schlechter 4052 (PRE, syn.!).

Agrostis eriantha Hack. var. planifolia Goossens &

Papendorf: 181 (1945), syn. nov. Type: South Africa,

[Gauteng], Irene, Doornkloof, Pole-Evans 666 (PRE,

holo.l).

ACKNOWLEDGEMENTS

The Botanical Education Trust is gratefully acknowl-

edged for the financial award that was granted to the

Taxonomic Problems in Plants of Conservation Impor-

tance Project. We would like to thank Mrs Lorraine

Mills of the Department of Nature Conservation, Gau-

teng, for fieldwork assistance, and Mrs Lyn Fish of

SANBI for assistance and advice throughout this project.

REFERENCES

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 Sur-

vey of South Africa No. 58. National Botanic Gardens/Botanical

Research Institute, Pretoria.

GOOSSEN, A.P. & PAPENDORF, M.C. 1945. The Genus Agrostis.

South African Journal of Science 4\ \ 181-184.

HACKEL, E. 1904. Vierteljahresschrift der Naturforschenden Gesells-

chaft in Zurich 49: 172.

RAIMONDO, D., VON STADEN, L., FODEN, W., VICTOR, J.E.,

HELME, N.A., TURNER, R.C., KAMUNDI, D.A. & MANYA-

MA, PA. (eds). 2009. Red List of South African plants. Strelitzia

25. South African National Biodiversity Institute, Pretoria.

VICTOR, J.E. 2006. Data Deficient flags for use in the Red List of

South African plants. Bothalia 36: 85-86.

J.E. VICTOR't, A.C. MASHAU' "and VJ. NGOBENF

* South African National Biodiversity Institute, Private Bag XlOl, 0001

Pretoria. E-mail: *j. victor @ sanbi.org.za / c.mashau@sanbi.org.za.

" Student affiliation: Department of Botany, University of Pretoria,

0002 Pretoria.

MS. received: 2012-01-13.

RUBIACEAE

TAXONOMIC NOTES ON SERICANTHE ANDONGENSIS MAD k'HE'N COMBINATION AND STATUS IN SERICANTHE FROM

LIMPOPO, SOUTH AFRICA

Rubiaceae Juss. is one of the five largest families

of flowering plants with over 13 000 species (Bremer

2009) and belongs in the order Gentianales Juss. ex

Bercht. & J.PresI (APG 111 2009; Reveal 2012b). Mem-

bers of Rubiaceae can be recognized in the vegetative

state by their opposite, sometimes whorled, entire leaves

and interpetiolar stipules with axillary colleters. The

flowers are usually bisexual or sometimes unisexual or

functionally unisexual and polysymmetric, often with

a narrow corolla tube and spreading lobes; the ovary is

inferior in most species, with a nectary or disc on top,

except in members of tribe Gaertnereae in subfamily

Rubioideae, which have a secondarily superior ovary

(Jansen et al. 1996), and the fruit is baccate, drupaceous,

or capsular (Stevens 200 (-[accessed December 2011]).

There is strong molecular support for three subfamilies:

Bothalia 42,2 (2012)

205

Cinchonoideae Raf., Dialypetalanthoideae Reveal

[replacing Ixoroideae Raf. (Reveal 2012a)] and Rubio-

ideae Bremek. ex Verde. (Bremer & Eriksson 2009).

The African genus Sericanthe Robbr. belongs in sub-

family Dialypetalanthoideae — tribe Coffeeae DC. (Rob-

brecht & Puff 1986; Relief & Leistner 2000). Currently

tribe Coffeeae includes the southern African genera Cof-

fea L., Sericanthe Robbr., Tricalysia A. Rich, ex DC.,

and the reinstated genus Empogona Hook.f (Bridson

2003; Davis et al. 2007; Tosh et al. 2009). The tribe is

characterized by very short stipules that are connate and

apiculate; axillary inflorescences paired at nodes; rela-

tively small flowers; distinctly 2-lobed styles with diver-

gent arms; small, few-seeded, fleshy fruits; and seeds

with a deep ventral groove (Relief & Leistner 2000;

Tosh et al. 2009).

Tricalysia was more than 40 years ago considered as

a very broadly circumscribed genus. Halle (1970) recog-

nized that several species were misplaced in Tricalysia

and removed them to the new genus Neorosea N. Halle.

Detailed revisionary work by Robbrecht (1978), how-

ever, showed that one of these species, N. jasminiflora

(Klotzsch) N. Halle {=Tricalysia jasminiflora (Klotzsch)

Benth. & Hook.f ex Hiem), was in fact a Tricalysia.

Unfortunately this species was the one Halle has chosen

as the type of Neorosea, and Neorosea therefore became

a synonym of Tricalysia, while the remaining species

were left without a generic name. Robbrecht (1978)

rectified this by publishing the new name Sericanthe.

Subsequently Tricalysia subgenus Empogona (Hook.f)

Robbr. (1979) was raised to genus level, defined by the

very short calyx tube with well-developed lobes, corolla

mostly bearded in the throat, and anthers with conspicu-

ously apical appendages (Tosh et al. 2009). The genus

Tricalysia, as currently defined, is one of the largest

genera of Rubiaceae in Africa and occurs in continen-

tal Africa (± 95 spp.), Madagascar (12 spp.) and the

Comoros (1 sp.) (Tosh et al. 2009).

Sericanthe comprises 21 named species, confined

to Sub-Saharan Africa (Klopper et al. 2006), with four

recently described from Lower Guinea (Sonke et al.

2012). Sericanthe is distinguished from Tricalysia and

Empogona by the presence of linear bacterial leaf galls

or nodules (Van Oevelen et al. 2001), wing-shaped col-

leters, pollen with a verrucate sexine (in contrast to

reticulate sexine in all other members of Coffeeae), 7-

to 9-merous flowers (4- to 7-merous flowers in Trica-

lysia and Empogona), and basifixed anthers (medifixed

in Tricalysia and Empogon) with enlarged connectives

(Tosh et al. 2009). The seeds of Sericanthe can also

be distinguished by a ± elliptic to circular hilar scar, ±

half the size of the seed, while Tricalysia has a shal-

low hilar groove (Bridson 2003) and Empogona a hilar

excavation (Bridson 2003: 468, as subgen. Empogona).

In Sericanthe the seed coat is smooth and shiny when

preserved in liquid but slightly striate when dry (Rob-

brecht 1978; Bridson 2003; tab. 98, B2 and Figure 2D),

whereas the seed surface of Tricalysia and Empogona

are reported to be reticulate or pusticulate (Retief &

Leistner 2000; Bridson 2003).

Bacterial leaf galls, or nodules, occur in only three

genera of Rubiaceae, namely in Psychotria (tribe Psy-

chotrieae of subfamily Rubioideae), Pavetta (tribe

Pavetteae of subfamily Dialypetalanthoideae) and Seri-

canthe (Van Oevelen et al. 2001), however the nodules

of Pavetta and Psychotria are usually punctate to ellip-

soid, rarely only shortly linear. These galls house endo-

symbiotic bacteria. The bacterial partners are strictly

host specific and have been identified in Pavetta, Psy-

chotria, and recently also in Sericanthe andongensis

(Hiem) Robbr. as Candidatus Burkholderia andongensis

and in S. petitii (N. Halle) Robbr. as Candidatus Burkhol-

deria petitii (Lemaire et al. 2011).

The only species of Sericanthe from South Africa

was formerly referred to Sericanthe andongensis (Hiem)

Robbr. var. andongensis (Robbrecht 1978; Coates Pal-

grave 2002; Retief 2003, 2006; Klopper et al. 2006),

following Robbrecht (1978, 1988) who recognized two

varieties of S. andongensis, namely var. andongensis

and var. mollis Robbr. These two varieties grow more

or less sympatrically and differ in hairiness on their leaf

undersurfaces (var. andongensis with scattered hairs and

var. mollis velutinous) and their leaf apices (var. andon-

gensis always has acuminate apices and var. mollis has

rounded, obtuse to acute or occasionally subacuminate

leaf apices). In her treatment of Sericanthe in Flora

zambesiaca, Bridson (2003) redefined Robbrecht’s

(1978, 1988) concept of var. andongensis and var. mollis

for the plants occurring south of the Zambezi River. The

varieties of subsp. andongensis grow ± sympatrically

to the north of the Zambezi River but are geographi-

cally separated to the south. In Bridson’s opinion there

are five distinct taxa in Zimbabwe and South Africa: the

non-montane element is treated as subsp. engleri, which

is confined to the granite outcrops and serpentine hills of

the Matobos, southwestern Zimbabwe, while the mon-

tane specimens have been grouped with other doubt-

fully named specimens as three provisional species (see

Sericanthe sp. A, sp. B, and sp. C). Bridson (2003: 512)

continued: ^Tricalysia legatii Hutch., from South Africa,

has not been included as a synonym of Sericanthe

andongensis var. andongensis because, in my opinion,

it is both taxonomically and geographically discrete’.

She goes on to provide a few morphological differences

between the South African taxon, Tricalysia legatii and

S. andongensis subsp. engleri (K.Krause) Bridson, the

geographically nearest taxon. Tricalysia legatii differs

from S. andongensis subsp. engleri in its ± glabrous,

often obovate leaves (subsp. engleri has pubescent stems

and leaves), and calyx limbs 6-9 mm long with an api-

culum 1-3 mm long (limb 4-6 mm long with apiculum

1-2 mm long in subsp. engleri).

In Bridson’s (2003) opinion, Sericanthe andongensis

subsp. andongensis var. andongensis is considered to

be confined to Angola, Democratic Republic of Congo

(DRC), and north of the Zambezi River in Malawi,

Mozambique, Tanzania, and Zambia. Neither S. andon-

gensis subsp. andongensis var. mollis nor S. andongen-

sis subsp. engleri occur in the Flora of southern A frica

region according to Bridson (2003), and our own inves-

tigation of tropical and southern African material of this

genus supports this opinion. Therefore S. andongensis

var. andongensis is a misapplied name for South African

material, leaving the South African taxon unplaced.

206

Bothalia 42,2 (2012)

Besides the distinct geographical separation between

S. andongensis subsp. andongensis (including both vari-

eties), subsp. engleri and Tricalysia legatii, leaf char-

acters such as hairiness, size, apex, and petiole length,

can be used to separate these entities. However, there

is great variation in the hairiness of leaves, a character

that might correlate with the different stages of develop-

ment of the leaves, since most plants are deciduous, with

the flowers appearing before the leaves. Differences in

hairiness of the leaves are not always a distinctive dis-

tinguishing character to separate these entities. On the

South African specimens of S. andongensis at Kew,

Robbrecht has frequently annotated the specimens ‘tend-

ing towards mollis' (i.e. var. mollis Robbr.) (Burrows,

pers. comm.). Although some South African plants do

seem to be pubescent when young, it would seem not

sufficiently so for Robbrecht to unequivocally place

them in var. mollis.

We accept Bridson’s opinion that Tricalysia legatii is

a distinct taxon on account of the differences mentioned

above, but the differences between it and the other sub-

species and varieties of S. andongensis are not sufficient

enough to elevate this taxon to species level. The pur-

pose of this paper is therefore to provide a subspecific

classification and combination in order to accommodate

the South African taxon and to provide a key to the cur-

rent accepted subspecies and varieties of S. andongen-

sis. The three unnamed, undescribed entities of Bridson

(2003), namely the montane element of S. andongensis

from Zimbabwe (sp. A, sp. B, sp. C), are not dealt with

in this paper.

All specimens cited are studied from herbarium

specimens housed in PRE. The most important type

specimens were seen on websites which are cited below.

Acronyms for herbaria are listed in Holmgren et al.

(1990).

TAXONOMY

Key to subspecies and varieties of Sericanthe andon-

gensis (modified from Bridson 2003)

la Mature stems usually not gnarled, with nodes and lateral

twigs widely spaced; leaves monomorphic or ± dimor-

phic, usually > 50 mm (up to 113 mm) long; leaf lamina

narrowly elliptic to broadly ovate, occasionally obovate;

petiole 5-10 mm long; Angola, DRC, Tanzania, Malawi,

Mozambique, Zambia (subsp. andongensis)'.

2a Leaves with only few scattered hairs below; leaf apex

mainly acuminate subsp. andongensis var. andongensis

2b Leaves velutinous below; leaf apex usually rounded,

obtuse to acute, very occasionally a few leaves with sub-

acuminate apices subsp. andongensis var. mollis

1 b Mature stems often gnarled, with nodes and lateral twigs

closely spaced; leaves clearly to markedly dimorphic,

usually < 50 mm long (sometimes young leaves up to 70

mm long); leaf lamina variously obovate, oblanceolate,

broadly elliptic to oblong; petiole up to 5 mm long; Zim-

babwe, South Africa:

3a Leaves softly pubescent below or occasionally with few

scattered hairs; leaf lamina rarely obovate; calyx limb

4-6 mm long; southwestern Zimbabwe subsp, engleri

3b Leaves glabrcsccnt below; leaf lamina often obovate;

calyx limb 6-9 mm long; South Africa: Limpopo ....

subsp. legatii

Sericanthe andongensis (Hiern) Rohhr. in Bulletin

dll .lardin Botanique de I’Etat a Bruxelles 48: 31 (1978).

Neorosa andongensis (Hiem) N. Halle: 270 (1970).

Type: Angola, Cuanza Norte, Pungo Andongo, on the

Calemba islands in the river Cuanza, Welwitsch 3133

(LISU, holo. — Aluka image!; BM — Aluka image!, K —

Aluka image!, LISC — ACTD image!, iso.).

Sericanthe andongensis subsp. andongensis var.

mollis Rohhr. in Bulletin du Jardin Botanique de I’Etat

a Bruxelles 48: 35 (1978). Type: DRC, without pre-

cise locality, Quarre 1437 (BR, holo. — Aluka image!;

SRGH — Aluka image!, iso.).

Sericanthe andongensis subsp. engleri (K.Krause)

Bridson in Flora Zambesiaca 5: 514 (2003). Type: Zim-

babwe, Matobo Dist., Kora Kora [Korakora] Stream

near Absent Farm, Plowes 1474 [BR, neotype — Aluka

image!, chosen by Robbrecht (1978); MO — digital

image!, P — digital image!, SRGH, isoneo.].

Sericanthe andongensis subsp. legatii (Hutch.) Jor-

daan & H.M.Steyn, stat. nov. Tricalysia legatii Hutch.,

A botanist in southern Africa: 306 (1946). Type: South

Africa, [Limpopo Province], Soutpansberg [Zoutpans-

berg], between Louis Trichardt and Farm Geluk, Sep-

tember 1908, Legat 23 (K, holo. — Aluka image!, PRE,

photo!).

Sericanthe andongensis subsp. andongensis var.

andongensis sensu Robbr.: 31 (1978), pro parte, as to

specimens cited from South Africa.

Semi-deciduous or deciduous shrub or tree up to 5 m

tall, usually much smaller, young twigs densely hairy

with spreading hairs, becoming glabrescent with age,

older stems covered with dark brown, flaking bark.

Stems straight, with conspicuous, closely spaced nodes,

knobbly due to leaf scars and stipule remains. Stipule

lohes sheathing at least at base, truncate to triangular,

1.5-2. 5 mm long, each bearing an awn, l-2(-3) mm

long, with silky hairs on outside and colleters inside.

Leaves opposite, papery to ± leathery, slightly glossy

above, pubescent or glabrescent below, occasionally

with insect galls (± rounded to ovoid hairy excres-

cences), lamina obovate, oblanceolate to elliptic, mark-

edly dimorphic, (20-)25-50(-70) x (10-)15-25(-45)

mm, apex obtuse to acute, base cuneate or sometimes

somewhat rounded, midrib hairy on both sides, promi-

nent below, lateral veins in 3^(-6) main pairs, promi-

nently raised below, tertiary reticulate venation obscure

to conspicuous; domatia inconspicuous or occasionally

as conspicuous hairy tufts; bacterial galls linear, usu-

ally situated along basal half of midrib; petiole l-3(-5)

mm long, pubescent. Flowers 8(9)-merous, solitary,

axillary, white, sweetly scented, usually before new

leaves appear, subsessile; bract and bracteoles, cupular,

1-3 mm long, each with 2 unequal pairs of awns. Calyx

limh 4-6( 8) mm long, closed in bud, with an apicu-

lum 1-2 mm long, splitting into 2 or 3 lobes at anthesis,

thin in texture and often translucent, silky haii^ outside.

Corolla salver-shaped, tube 7-13 mm long, silky hairy

outside except at base; lobes 7-12 mm long and slightly

twisted, ± truncate with acuminate tip but acumen not

central in mature lobes. Stamens arising at throat, fila-

ments up to 1 mm long, glabrous, anthers 4-6 mm long,

exserted. Ovaty 2-locular, ovules 2 per locule, style gla-

brous, 2-lobed, reaching or slightly exceeding anthers.

Bothalia 42,2 (2012)

207

Fruit a spherical drupe, ± 10 mm diam., orange or red. Etymology. Sericanthe is derived from the Greek

crowned by persistent calyx. Seeds either 1 per chamber words serikos = silk and anthos = flower, referring to

and hemispherical or 2 per chamber and quadriform, ± the silky hair-coat of the flower parts (Glen 2004). The

3^ mm in diam., dark brown, striate. Figures 1 & 2. specific epithet honours Charles Edward Legal (1876-

FIGURE 1. — Sericanthe andongensis subsp. legatii: A, flowering twig; B, stipule; C, flower. A-C, from Galpin 9557 (PRE). Scale bar: 10 mm.

Artist: Marietjie Steyn.

208

Bothalia 42,2 (2012)

FIGURE 2. — Sericanthe andongen-

sis subsp. legalii: A, fruiting

twig; B, leaf with bacterial

galls along midrib; C, fruit;

D, half-spherical seed with

striate coat. A, C, & D, from

Stalmam 1466 (PRE); B,

from McMurtry 10271 (PRE).

Scale bar: 10 mm. Artist:

Marietjie Steyn.

1966), who was bom in Scotland and came to South

Africa in 1 898 as a forester in the Cape Forestry Depart-

ment. He was later transferred to the former Transvaal

were he was Chief Conservator and Chief Conservator

of Forests for South Africa from 1913 to 1931, after

which he returned to England (Glen & Germishuizen

2010). Legal collected the type material in the Soutpans-

berg, near Louis Trichardt in 1908, from which Hutch-

inson in 1 946 described the species. Sericanthe belongs

to the same tribe as Cqffea, namely tribe Coffeeae, and

occurs in the northern parts of the Limpopo Province,

hence the common name Venda-coffee or silky-coffee as

in Van Wyk et at. (2011) under Tricalysia legafii.

Diagnostic characters: Sericanthe andongensis

subsp. legatii differs from subsp. andongensis, occur-

ing from Angola to Tanzania, and subsp. engleri from

the Matopos in Zimbabwe in having longer calyx limbs,

usually longer than 6 mm. For more differences see key.

Distribution and habitat: Sericanthe andongensis

subsp. legatii is confined to the Soutpansberg, Blou-

bcrg, and Wolkberg regions of Limpopo (Figure 3), thus

endemic to the Soutpansberg and Wolkberg Centres of

Endemism (Van Wyk & Smith 2001). It is recorded to

grow in bushveld, open woodland, or forest margins on

mountain or hill slopes, and rocky outcrops, often on

quartzite or sandstone.

Selected specimens

LIMPOPO. — 2229 (Waterpoort): Zoutpansberg Dist., Farm Bud-

worth, (~BB), Aug. 1958, Mogg 28281\ Zoutpansberg Dist., Goro, (-

CD), Jul. 1996, Barker 3; Louis Trichardt, Mountain Inn, (-DD), May

1966, Theron 1489. 2230 (Messina): Louis Trichardt Dist., ± 15 km

ENE of Louis Trichardt on the Witviag road. Farm Studholme, (-CC),

Apr. 1997, Burgoyne 6181\ SE of Nwanedi Reserve, N of Gundani Vil-

lage, just W of the Brachystegia spiciformis forest, (-DA), Mar. 2002,

McMiirliy 10271; Venda, Tshaulu Mission, Mutanzhela, (-DC), April

1980, Van Wyk 3812. 2328 (Baltimore): Polokwane [Pietersburg] Dist.,

Blouberg, Fami Leipsig, (-BB), Apr. 1961, Strey <& Schlieben 8470.

2329 (Pietersburg): Polokwane [Pietersburg] Dist., eastern foothills of

Blauwberg, (-AA), May 1953, Codd 7937; Zoutpansberg Dist., Fanu

‘Llewellyn’ 35 LS, hill opposite house, (-AB), Jul. 1981, Venter 6196;

Hanglip, near Louis Trichardt, (-BB), Dec. 1945, Gerstner 5760. 2330

(Tzaneen): near Louis Trichardt, Entabeni Forest Station, (-AA), Sept.

1929, Galpin 9557; Spelonken Dist.; Tshakhuma, (-AB), Nov. 1931,

Ohermeyer TRV30580; Daviesville, Setali Mission, (-AC), May 1945,

Gerstner 5484; Letaba Dist., Modjadji’s Reserve, near Duiwelskloof,

(-CB), Apr. 1937, Krige 104. 2430 (Pilgrim’s Rest): Lekgalameetse

Nature Reserve, Haffenden Heights, (-AA), Jan. 1978, Stalmans 1466;

Letaba Dist., Farm Cyprus, (-AB), Nov. 1960, Rcnny 139.

Bothalia 42,2 (2012)

209

FIGURE 3. — Known distribution of Sericanthe andongensis subsp.

legatii.

ACKNOWLEDGEMENTS

We would like to thank the artist Marietjie Steyn for

the line drawings and John and Sandie Burrows for stud-

ying the type material at Kew and for useful comments

on this manuscript. We also want to thank two anony-

mous reviewers for their suggestions.

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210

Bothalia 42,2 (2012)

IRIDACEAE

NOTES ON MORAEA SUBG. VISCIRAMOSA (IRIDOIDEAE), INCLUDING THE NEW SPECIES MORAEA SALDANHENSIS VKOU THE

ATLANTIC COAST OF SOUTH AFRICA, RECOGNITION OF M INCONSPICUA SUBSP. NAMAQUENSIS, AND POLLINATION

BIOLOGY IN M RIVULICOLA

Novelties in the largely sub-Saharan genus Moraea

Mill. (Iridaceae: Iridoideae), now comprising over

215 species (Goldblatt & Manning 2009), are still

being regularly discovered in southern Africa. Florally

diverse, Moraea is recognized in Iridoideae by a bifacial

and channelled (rarely terete or plane) leaf blade, and

conns of a single intemode derived from a lateral bud.

Most species have /ra-like flowers with flattened, peta-

loid style branches to which the stamens are appressed

(Goldblatt 1986), but the occurrence of a variety of other

floral types (Goldblatt 1998) makes exact floral defini-

tion of the genus impractical, although most species

have free tepals and partially to completely united fila-

ments.

One of the more distinctive groups in the genus is

subg. Visciramosa Goldblatt, distinguished by its well-

branched stems sticky below the nodes. Moraea saldan-

hensis, the new species described here, a recently dis-

covered local endemic of sandy coastal flats and hills

north and east of Saldanha Bay in Western Cape, is

unusual in subg. Visciramosa (now with 8 species) in its

slate blue-grey tepals with off-white markings, and with

the claws longer than the limbs. Other species in the

subgenus have yellow to buff or brown flowers although

M. viscaria (L.f ) Ker Gawl. is characterized by its white

flowers. We report the existence of a population of M.

hituminosa (L.f.) Ker Gawl. with blue-mauve tepals, as

well as white-flowered populations of M. elsiae Gold-

blatt and M. inconspicua Goldblatt, both until now

recorded as having yellow or, in M. inconspicua, also

buff to brown flowers. Namaqualand populations of the

widespread winter rainfall species M. inconspicua are

distinctive in their coiled leaves and paler flowers, and

we distinguish them here as subsp. namaquensis. The

current key to the subgenus (Goldblatt 1986) included

only five species and was already outdated by the rec-

ognition of M. vespertina Goldblatt & J.C. Manning in

2000 and M. simplex Goldblatt & J.C. Manning in 2004

(Goldblatt & Manning 2000, 2004). We provide a new

key to all eight species and one subspecies now recog-

nised in subgen. Visciramosa.

In addition, we report an unusual pollination sys-

tem in Moraea rivulicola Goldblatt & J.C. Manning

(subg. Vieusseuxia). Pollination strategies in Moraea

are diverse, and include large-bodied anthophorine and

honey bees foraging for nectar, female bees of several

families foraging for pollen, hopbine beetles, muscid,

flesh and blow flies, and eumenine wasps (in one species

to date: Goldblatt et al. 2005). In M. rivulicola pollen

transfer is accomplished by species of eumenine wasps

in a system that closely matches that reported in three

species of Ferraria (Goldblatt et al. 2009), which have

similar floral odours and nectar characteristics.

Wc examined all relevant collections at BOL, NBG,

PRE, and SAM, the primary southern African herbaria

(acronyms after Holmgren et al. 1990). The distribution

of M inconspicua subsp. inconspicua (Figure 2) is based

on this material although collections for this taxon are

not cited.

New taxa

1. Moraea saldanhensis Goldblatt & J.C. Manning,

sp. nov.

TYPE. — Western Cape. 3217 (Vredenburg): rocky

coastal sandveld near Jacobsbaai at Mauritzbaai, ( DD),

20 Oct. 2011, Manning 3310 (NBG, holo.; MO, iso.).

Plants 150-350 mm high. Corm obconic-subglobose,

25^0 mm diam., bearing numerous small cormlets at

base; tunics pale brown, becoming fibrous, with strong

vertical ribs joined by fine fibres in herringbone pattern.

Stem ± erect, firm, sticky below nodes, usually branch-

ing at all nodes, with 2, rarely 3, branches per node;

branches sharply flexed outward above sheaths and then

± horizontal, flexed upward at apex. Foliage leaves 2,

lowermost inserted below ground, linear, channelled,

lowermost longest, initially suberect, trailing above, up

to 400 mm long, 4-6 mm wide when opened flat, sec-

ond leaf somewhat shorter; sheathing leaves up to 28

mm long, distal 5-6 mm dry and brown, ± truncate or

subacute. Rhipidia several-flowered; spathes green, api-

ces subacute to ± truncate, purple near tips and turning

dry and brown with age in upper 5 mm, inner 22-25 mm

long, outer ± 1/2 to 2/3 as long, completely sheathing.

Flowers slate blue-grey to purple, outer tepals coppery

pink outside and with off-white nectar guides minutely

speckled with blue dots, claws whitish, minutely speck-

led grey-blue, nectar guide at base of outer tepal limbs

dark blue-grey; tepals free, smooth, outer ± 18 x 5. 5-7.0

mm, initially spreading, later half-reflexed sometimes

recurving distally, claws narrow, ± 10 mm long, thus

slightly exceeding limbs, suberect; inner tepals up to

14 X 3 mrn, oblanceolate, limbs spreading horizontally

or later half reflexed. Stamens with filaments ± 8 mm

long, free but closely adhering in smooth, cylindric col-

umn ± 6.5 mm long, free and diverging distally; anthers

3-4 mm long, appressed to style branches, dark purple,

pollen orange-red. Ovajy ± 8 mm long, usually included

in spathes; style dividing at apex of filament column,

branches ±6x2 mm, sharply diverging and extended

horizontally, white, flushed purple distally; stigma lobe

transverse, with prominent central cusp ± 2 mm long,

acute or bifid at apex, crests arching upward distally

and 4 5 mm long, purple, intensely so in distal half

Capsules and seeds unknown. Chromosome number

unknown. Flowering time: late Sept.-Nov. Figure 1.

Distribution and ecology: Moraea saldanhensis is

currently known from two sites in sandy ground among

granite rocks along the coast of Western Cape, one

between Jacobsbaai and Saldanha and a second near

Vredenburg (Figure 2). Unusually late flowering in the

area, it blooms after most of the spring flora, and the

Bothalia 42,2 (2012)

211

FIGURE 1 . — Moraea saldanhensis, Goldblatt & Manning 13405 (NBG). A, plant; B, outer tepal, with detail of nectary; C, inner tepal; D, stamens

and style; E, detached style branch. Scale bar: A, 10 mm; B-E, 2 mm. Artist: John Manning.

slate-blue flowers are inconspicuous among the drab

green or dry vegetation. The first record was made by

the Cape botanist, N.A. Helme, in November 2006, and

two years later the Jacobsbaai site was discovered by

local resident Koos Claasens, who helped us make the

type collection in October 2010. The narrow distribution

of the species, in an area undergoing rapid development

along the coast and inland, leaves it in a parlous conser-

212

Bothalia 42,2 (2012)

FIGURE 2. — Distribution of Moraea saldanhensis. A; M inconspicua

subsp, inconspicua, o; M. inconspicua subsp. namaquensis, •,

vation state and it must be regarded as vulnerable (VU)

if not endangered (EN) using the definitions and termi-

nology of Raimondo et ah (2009).

Diagnosis and relationships', clearly a member of

subgen. Visciramosa Goldblatt of Moraea (Goldblatt

1976a), M. saldanhensis has the large obconic corms

with brown tunics, stems sticky below the nodes, rela-

tively blunt rhipidial spathes and multi-branched stems

characteristic of the alliance. Vegetatively, the relatively

short spathes recall M. elsiae and M. inconspicua and

it is with the latter that M. saldanhensis is most easily

confused. Both have small flowers with well-developed

style branches and prominent style crests. The flowers of

M. inconspicua are yellow or partly to largely brown or

occasionally white, and the tepal limbs of both whorls

are half to fully reflexed (Goldblatt 1986). The larger

outer tepals of M. inconspicua are 12-18 mm long with

undulate to ± plane limbs 9-12 x 6-7(8) mm and claws

4-7 mm, thus ± one third to half as long as the limbs

(Figure 3; Table 1 ). This stands in marked contrast to the

flowers of M. saldanhensis, which has outer tepals ± 1 8

mm long initially spreading, later half-reflexed and dis-

tally recurving, and narrow erect claws ±10 mm long,

slightly longer than the limbs, the opposite of the situ-

ation in M inconspicua. Moraea saldanhensis also has

a slightly longer filament column, ± 6.5 mm long (vs.

3. 5-6.0 mm in southern populations of M. inconspicua,

i.e. subsp. inconspicua). Pollen in M saldanhensis is

orange-red and the ovary is included whereas in M

inconspicua the ovary is usually exserted and the pollen

is typically yellow, although populations with red pollen

are known, notably among the Namaqualand popula-

tions, segregated as subsp. namaquensis (see below).

Additional specimen

WESTERN CAPE. — 3217 (Saldanha): western edge of Vreden-

burg, just S of road to De Klip, Saldanha Granite Strandveld, (-DD), 5

Oct. 2006, Heline 4162 (NBG, photo).

2. Namaqualand populations of Moraea inconspicua

are immediately recognized by their tightly coiled

leaves, dull cream-coloured, ivory to buff or brown

flowers with the tepal limbs usually reflexed to lie in a

vertical plane, and by their bright orange pollen. Plants

from south of Namaqualand have channelled, ascend-

ing to trailing leaves and usually smaller flowers (Table

2), mostly bright yellow to buff, sometimes brown or

rarely white, with the tepal limbs either fully reflexed

or spreading to lie ± 45° from the horizontal, and usu-

ally yellow, rarely red pollen. The Namaqualand plants

typically grow with their corms tightly wedged among

rocks whereas plants from the south occupy a diverse

range of habitats but mostly open stony or sandy flats.

The combination of morphological and geographical

differentiation are consistent with the recognition of

the Namaqualand populations as a distinct subspecies,

and with similar taxonomic treatments that we have

applied to races of Freesia viridis (Aiton) Goldblatt &

J.C. Manning (Manning & Goldblatt 2010) and Tritonia

securigera (Aiton) Ker Gawl. (Goldblatt & Manning

2006).

Moraea inconspicua subsp. namaquensis Goldblatt

& J.C. Manning, subsp. nov.

TYPE. — Northern Cape. 2917 (Springbok): southeast

of Driekoppie farmhouse, (-DB), 3 Oct. 1981, Van Ber-

kel 439 (NBG, holo.; MO, iso.)

Like subsp. inconspicua but foliage leaves coiled

with surface plane, 2-3 mm wide. Rhipidia with outer

spathes mostly 20-28 mm long. Flowers ivory, creamy

yellow, creamy pink, pale yellow or brown; tepals usu-

ally reflexed to lie vertically, outer 15-18 mm long,

claws 5-7 mm long. Stamens with filaments 6-8 mm

long; anthers 3^ mm long, pollen usually red. Style

branches 5-6 mm long, crests 3-4 mm long. Flowering

time'. mid-Sept.-late Nov.

Distribution and ecology: restricted to the higher-

lying edge of the western escarpment in Namaqualand,

between the Kamiesberg and Steinkopf (Figure 2).

Diagnosis', distinguished from typical forms by the

coiled leaves with flat blades up to 3 mm wide. Plants

of subsp. inconspicua, which occurs south of Namaqua-

land, from the Bokkeveld Mtns to Port Elizabeth, have

channelled leaves that are initially erect later trailing

but never coiled. Other vegetative and floral features

differ only quantitatively, with considerable overlap

but on average subsp. namaquensis has slightly larger

rhipidial spathes and floral features than the typical

TABLE 1. — Selected features of Moraea inconspicua subsp. inconspicua, subsp. namaquensis, and M. saldanhensis.

Bothalia42,2 (2012)

213

TABLE 2. — Selected floral characteristics of typical, yellow-flowered Moraea hHuminosa and the mauve blue-flowered morph. Only living plants

or well-pressed, fully open flowers were measured.

subspecies. In subsp. namaqiiensis the outer rhipidial

spathes are 20-28 mm long, outer tepals 15-18 mm

long, and anthers 3-4 mm long compared with rhip-

idial spathes 15-26 mm long, outer tepals 12-15 mm

long and anthers 2. 0-3. 3 mm long in subsp. inconspicua

(Table 1). M. inconspicua has not, to date, been recorded

between the Bokkeveld Mtns and the Kamiesberg, an

unexpected gap in its range that evidently represents the

geographical disjunction separating the two subspecies.

Additional specimens

NORTHERN CAPE.— 2917 (Springbok): Steinkopf, (-BC), Oct.

1934. Herre s.n. (BOL, PRE); ± 5 km W of Steinkopf, (-BC), 10 Oct.

1987, Williamson 3776 (NBG); near top of Spektakel Pass, (-DA), 11

Sept 1993, Goldblatt & Manning 9714 (MO, NBG); 7 km S of Spring-

bok on Camping Ground, (-DB), 12 Oct. 1989, Greater 21661 (PRE);

hills E of Nababeep, (-DB), 14 Oct. 1974, Goldblatt 3054 (MO, NBG,

PRE). 3018 (Kamiesberg): 2 km E of Kamieskroon, (-AA), 28 Sept.

1976, Goldblatt 4255 (MO, NBG); Kamiesberg, Farm Damsiand, (-

AC), 19 Oct. 2007, Snijman 2206 (NBG).

Notes on divergent flower colour morphs in other

species of subgen. Visciramosa

1 . Moraea bitiiminosa: with pale to deep yellow flow-

ers over most of its range, which stretches from Tulbagh

and the Cape Peninsula eastward to the Bredasdorp

Mtns, M. bituminosa has the largest flowers in subgen.

Visciramosa, with tepals up to 30 mm long and open

flowers up to 40 mm in diameter (Goldblatt 1976b).

Unusual for the species and subgenus, a population of

M. bituminosa from Jonkershoek has blue-mauve flow-

ers. In November 2011 we found a second site for the

blue-mauve-flowered morph of the species and we were

able to investigate these alive. The outer tepals have nec-

tar guides consisting of a band of fine black dots at the

limb bases edged with a broad white band, the anthers

are dark purple-black and the pollen is bright red. Yel-

low-flowered plants have nectar guides consisting of a

patch of black spots on a dark yellow background and

invariably yellow anthers and pollen. The blue-mauve

flowers closely match those of yellow-flowered plants

except that the filament column is mostly longer, the

tepal claws slightly longer, and the style crests signifi-

cantly longer than the typical yellow morph (Table 2).

We are uncertain about the significance of the blue

mauve-flowered morph of Moraea bituminosa and

assume that the Jonkershoek and Tulbagh Valley plants

are the same genetic race. The morph is embedded

within the range of the yellow-flowered morph, and at

the Tulbagh valley site yellow-flowered plants grew

adjacent to those with blue-mauve flowers (which at

this locality had smaller flowers) with no evidence of

intermediates. At this juncture we merely report the bio-

logical situation. The sympatry in the Tulbagh Valley of

yellow and blue-mauve-flowered morphs raises the pos-

sibility that the latter may be a separate species.

2. Moraea elsiae: with relatively small flowers, M.

elsiae stands out in subgen. Visciramosa in having the

style branches much reduced in size and lacking style

crests, and narrower than the subtending anthers, the tips

of which exceed the style branches (Goldblatt 1976b;

1986). The tepals have until now been reported as yel-

low and the limbs of both whorls spread slightly below

the horizontal; anthers and pollen are also yellow. Plants

from near Elim and Kleinmond, however, stand out in

having white tepals, the outer ± 16 x 5-6 mm, somewhat

smaller than elsewhere in the species, and with anthers

2. 5-3.0 mm long. In yellow-flowered plants the outer

tepals are 1 5-22 x 9 mm and anthers are 3^ mm long.

The recorded range of Moraea elsiae is from near

Mamre and the Cape Peninsula to Still Bay (Goldblatt

& Manning 2009). The white flowered populations thus

fall within the range of the species and are evidently a

distinct regional race. At one locality white-flowered

M. elsiae grew intermixed with plants of M. incon-

spicua but flowers opened in the mid- to late afternoon,

whereas those of M. inconspicua opened in the morning

and wilted at about the time those of M. elsiae opened.

White-flowered specimens

WESTERN CAPE.— 3419 (Caledon): Kleinmond, (-AC), 26 Nov.

1949, de Vos 1531 (NBG); clay slope ± 4 km W of Elim, (-DA), 9

Nov. 2011, Goldblatt & Porter 13740 (MO, NBG).

3. Moraea inconspicua: as circumscribed at present,

M. inconspicua is widespread in the southern Afri-

can winter rainfall zone, extending from northern

Namaqualand to Port Elizabeth (see subsp. namaqiiensis

described above). Flowers in the species are variable in

colour but mostly yellowish to brown, buff, occasion-

ally ivory, and pollen is yellow or reddish. Plants from

Elim, west of Bredasdorp, that have come to our atten-

tion, have white tepals and the anthers are ± 2.5 mm

long, the style branches ± 3 mm long, and the style

crests ± 1 .5 mm long, thus as small or smaller than pre-

viously recorded for the species (anthers 2^ mm, style

branches mostly 4-5 mm, style crests 3^ mm long).

The pollen is red. The white-flowered populations from

the near Elim fall within the geographic range of yel-

low to brown-flowered plants and do not in our opinion

merit taxonomic recognition although they evidently

represent a distinct race of the species, distinguished by

tepal colour and in the smaller size of the anthers, style

branches and crests and red pollen. Superficially similar

to white-flowered M. viscaria, these populations are dis-

tinguished by the reduced size of the style crests, which

in M. viscaria are well developed and at least half as

long as the branches.

214

Bothalia 42,2 (2012)

FIGURE 3. — Moraea inconspicua

subsp. inconspicua, Bo-

Hermon, Elandsberg Farm,

without voucher. A, flower; B,

outer tepal; C, inner tepal; D,

stamens and style branches; E,

detached style branch. Scale

bar: 2 mm. Artist: John Man-

ning.

White-flowered specimen

WESTERN CAPE. — 3419 (Caledon): clay slope ± 4 km W of

Elim, (-DA), 9 Nov. 2011, Goldblalt & Porter 13743 (MO, NBG).

Key to taxa of subg. Visciramosa

1 . Style crests absent or rudimentary:

2. Style branches plane and appressed to opposed anther; flow-

ers yellow or white M. elsiae

2. ’ Style branches filiform, terete, extending between stamens;

flowers pale yellow M simplex

1.’ Style crests moderately to well developed, at least half as

long as style branches and usually > 2 mm long:

3. Flowers relatively small, outer tepals 13-23 mm long;

anthers 2. 5-4.0 mm long; inner rhipidial spathes usu-

ally 1 8-25 mm long:

4. Flowers slate blue; outer tepal limbs ± 8 mm long, slightly

shorter than claws M saldanhensis

4. ’ Flowers white, yellow, buff or ± brown; outer tepal limbs

9-15 mm long, usually ± twice as long as claws:

5. Flowers white, sweetly scented and opening mid- to late

afternoon; tepals 16-23 mm long with limbs laxly spread-

ing M viscaria

5. ’ Flowers usually yellow, buff or ± brown, occasionally white,

not noticeably scented or with spicy odour and opening

mid- to late morning; tepals 13-18 mm long with limbs

spreading to reflexed M. inconspicua

6. Leaves trailing, conduplicate; tepals brown to yellow, rarely

white; plants from Cape Floristic Region .... subsp. inconspicua

6. ’ Leaves coiled, ± flat; tepals pale whitish to buff or brown;

plants from Namaqualand subsp. namaquensis

3.’ Flowers large, outer tepals (22-)26-40 long; anthers at least

4 mm long; inner rhipidial spathes (23-)25^0(-55) mm long:

7. Foliage leaves usually 4-6, lower 3-5 basal; flowers white,

sometimes fading blue-grey, or pale slate blue and open-

ing at dusk M. vespertina

7. ’ Foliage leaves 2 or 3, lower I or 2 basal; flowers yellow,

blue-mauve or light brown to buff with yellow nectar

guides and opening at or before midday:

8. Flowers bright yellow with yellow anthers and pollen, or

bluc-mauve with blackish anthers and red pollen; only

outer tepals with nectar guides M. hituminosa

8.’ Flowers predominantly buff to light brown with light brown

anthers and pollen; inner and outer tepals with yellow nec-

tar guides at base of limbs M. bubaUna

Pollination «/ Moraea rivulicola

Endemic to central and northern Namaqualand, M.

rivulicola Goldblatt & J.C. Manning (subgcn. Vieus-

seiixia) is restricted to locally wet sites along streams,

seeps and the edges of seasonal marshes. Among species

of subgen. Vieiissieuxia it is distinguished by a relatively

robust habit, taller plants reaching over 1 m, compara-

tively large, dull coloured flowers with outer tepals dull

cream to pale yellow-green, sometimes flushed with

pink and the trilobed inner tepals and style branches

are light brown. Plants flower from late August through

September to early November at higher elevations. At a

site in the higher Kamiesberg, central Namaqualand, on

the Farm Karas, plants were in flower in the last days of

October {Goldblatt & Porter 13683 , MO, NBG) when

by chance, we had the opportunity to observe pollinator

activity.

Flowers open in mid-morning, last three days, and

during the day have an unusual, unpleasant scent, a sour,

aminoid odour mixed with faint notes of molasses. From

10:00 to 12:00 flowers were actively visited by mode-

rate-sized, black eumenine wasps, Allepipona sp. and

Knemodynerus sp. Both species of wasp were frequent

visitors, individuals entering the flowers by crawling

down the channel formed by the claw of an outer tepal

and closely opposed style branch. When individuals

emerged white pollen could clearly be seen in the dor-

sum. We infer that the wasps were foraging for nectar

contained at the base of the outer tepal limbs. Nectar is

secreted from shiny nectaries both at the limb base and

on the abaxial surface of smalt ridges of tissue at the

limb bases. When examined with a hand lens the nectary

surfaces of freshly opened flowers carried minute spots

of liquid (as glistening spots). Overnight the tepal bases

accumulated a large droplet of fluid, which when tasted

on the tongue had no trace of sweet flavour.

Pollination by eumenine and masarine wasps (Eume-

nidae: Eumeninae and Masarinae) in Iridaceae is known

in just one species of Moraea and three species of the

related genus, Ferraria (Gess & Gess 1989; Goldblatt

et al. 2009; Goldblatt & Manning 2011). The wasp pol-

linated Ferraria species, F. divaricata Sweet, F macro-

chlamys (Baker) Goldblatt & J.C. Manning, and F. vari-

ahilis Goldblatt & J.C. Manning, also have dull-coloured

flowers, unusual, aminoid-type odours, and produce

watery nectar of low sugar concentrations (usually less

than 10% sucrose equivalents). Wasp species involved

in pollination of these Ferraria species included the

Bothalia 42,2 (2012)

215

eumenines Delta coffer, Delta sp. and Allepipona ery-

throspina and the masarines Celonites capensis and

Jugurtia koeroegabensis. Wasp pollination very simi-

lar to that in species of Ferraria has evidently evolved

independently in Moraea riviilicola. This is the second

example of eumenine wasp pollination in Moraea. Two

species of Tricarinodynerus and one of Paracliilus were

reported by Goldblatt et al. (2005) as pollinators of M.

inconspicua Goldblatt (subgen. Visciramosa) in northern

Namaqualand.

ACKNOWLEDGEMENTS

We thank Koos Claasens of Jacobsbaai and Nick

Helme of Cape Town for bringing the new Moraea

saldanhensis to our attention and for their help in col-

lecting the species, and Elizabeth Parker and Len Por-

ter for assistance in the field. Field work was supported

in part by a grant from the Mellon Foundation. We also

acknowledge the help of Simon van Noort, Iziko Mu-

seums of Cape Town, and James M. Carpenter, Ameri-

can Museum of Natural History, for providing identifica-

tions for the wasps captured on Moraea riviilicola. Wasp

specimens are deposited at the South African Museum,

Cape Town.

REFERENCES

GESS, S.K. & GESS. F.W. 1989. Flower visiting by masarid wasps in

southern Africa (Hymenoptera: Vespoidea: Masaridae). Annals

of the Cape Provincial Museums. Natural Histoiy 1 8: 95-1 34.

GOLDBLATT, R 1 976a. Evolution, cytology and subgeneric classifica-

tion in Moraea (Iridaceae). Annals of the Missouri Botanical Gar-

den 63: 1-23.

GOLDBLATT, P. 1976b. The genus Moraea in the winter rainfall

region of southern Africa. Annals of the Missouri Botanical Gar-

den 63:657-786.

GOLDBLATT, P. 1986. The moraeas of southern Africa. Annals of

Kirstenbosch Botanical Garden 14: 1-224.

GOLDBLATT, P. 1998. Reduction of Barnardiella, Galaxia, Gynan-

driris, Hexaglottis, Homeria and Roggevetdia in Moraea (Iri-

daceae: Irideae). Novon 8: 371-377.

GOLDBLATT, R, BERNHARDT, P. & MANNING, J.C. 2005. Pol-

lination mechanisms in the African genus Moraea (Iridaceae:

Iridoideae): floral divergence and adaptation for pollen vector

variability. Adansonia 27: 21-46.

GOLDBLATT, R, BERNHARDT, R. & MANNING, J.C. 2009. Adap-

tive radiation of the putrid perianth: Ferraria (Iridaceae: Irideae)

and its unusual pollinators. Plant Systematics and Evolution 278:

53-65.

GOLDBLATT, P. & MANNING, J.C. 2000. New species of Moraea

(Iridaceae-Iridoideae) from southern Africa. Novon: 10: 14-22.

GOLDBLATT, P. & MANNING, J.C. 2004. New species of Aw (Cro-

coideae) and Moraea (Iridoideae), and taxonomic notes on some

other Alrican Iridaceae. Novon 14: 288-298.

GOLDBLATT, P. & MANNING, J.C. 2006. Notes on the systematics

and nomenclature of Tritonia (Iridaceae: Crocoideae). Bothalia

36: 57-61.

GOLDBLATT, P. & MANNING, J.C. 2009. New species of Moraea

(Iridaceae: Iridoideae), with range extensions and miscellaneous

notes for southern African species. Bothalia 39: 1-10.

GOLDBLATT, P. & MANNING, J.C. 2011. Systematics and biology

of the African genus Ferraria (Iridaceae: Irideae). Bothalia 41:

1^0.

HOLMGREN, P.K., HOLMGREN, N.H. & BARNETT, L.C. 1990.

Index Herbariorum. Part. I: The Herbaria of the World. New

York Botanical Garden, New York.

MANNING, J.C. GOLDBLATT, P. 2010. Botany and horticulture of the

genus Freesia (Iridaceae). Strelitzia 27. South African National

Biodiversity Institute.

RAIMONDO, D., VON STADEN, L., FODEN, W., VICTOR. J.E.,

HELME, N.A., TURNER, R.C., KAMUNDI, D.A. & MANYA-

MA, RA. (eds). 2009. Red List of South African Plants. Strelitzia

25.

P. GOLDBLATT’ and J.C. MANNING"

'B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.

O. Box 299, St. Louis, Missouri 63166, USA.

" Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town. / Research Centre for

Plant Growth and Development, School of Biological and Conserva-

tion Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private

Bag XOl, Scottsville 3209, South Africa. E-mail: J.Manning(g)sanbi.

org.za.

MS. received: 2011-12-21.

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Bothalia 42,2: 217-245 (2012)

Nomenclature and typification of southern African species of

Euphorbia

P.V. BRUYNS*

Keywords: Euphorbia, Euphorbiaceae, lectotypes, neotypes, typification

ABSTRACT

Types have been located for most of the 1 85 species of Euphorbia L. that are known to occur naturally in southern Africa

and also for most of their synonyms. Lectotypes or neotypes are selected where possible for those names for which a holotype

cannot be found. The synonymy largely follows previous accounts and reasons are given where new synonymy is proposed.

Euphorbia hiittonae N.E.Br. is reinstated at the level of species and E. franksiae var. zuluensis A. C. White et at. is raised to

the level of species as E. gerstneriana Bruyns, nom. nov. A new name, E. radyeri Bmyns, is provided for the rhizomatous

plants previously referred to as E. caerulescens Haw., which is synonymous with E. ledienii A. Berger.

INTRODUCTION

The large, cosmopolitan genus Euphorbia L. is rep-

resented in southern Africa (taken here to include the

countries Botswana, Lesotho, Namibia, South Africa,

and Swaziland) by 185 species. For these there are 368

validly described names at the level of species. In this

listing only naturally occurring species are included.

These are distributed among the four subgenera estab-

lished by Bruyns et al. (2006) and based on molecu-

lar and morphological data as shown in Table 1, where

additional information on growth habit is also supplied.

Many of the succulent species such as E. globosa Sims,

E. meloformis Aiton and E. obesa Hook.f are popu-

lar subjects in specialist collections around the world,

but the other species have little economical use, though

their medicinal use is probably underestimated. The

last revisions of the genus for this region were those of

N.E. Brown (1911-1912, 1915), with the succulent spe-

cies receiving further attention by White et al. (1941).

The taxonomy of Euphorbia in southern Africa remains

disorganised, with many names applied in different her-

baria in South Africa to quite different species. In an

attempt to bring order to the situation, types have been

located and, where these are missing, lectotypes or neo-

types are selected as applicable.

MATERIALS AND METHODS

For each validly published name for a southern

African species of Euphorbia, the protologue was con-

sulted in the relevant literamre. Type specimens have

been searched for among material in the herbaria B,

BM, BOL, G, GRA, K, KMG, M, NBG, NH, NU, NY,

OXF, P, PRE, S, SAM, SBT, W, WIND, WRSL, WU,

Z (herbarium acronyms according to Holmgren et al.

(1990)). A specimen is taken as the holotype if it was

indicated as such by the author, or if it is clear from

where it is located relative to where the author worked

that it must be the holotype. In many cases it has proved

to be impossible to be sure which specimen is the holo-

* Bolus Herbarium, University of Cape Town, 7701

MS. received: 2011-10-18

type. In such instances, a lectotype is generally selected

from among the duplicates of the ‘type number’ or from

among the ‘syntypes’ mentioned by the author in the

protologue. A particular specimen is chosen over oth-

ers according to its quality or, in the case of syntypes,

according also to how widely duplicates (if any) are rep-

resented. In cases where no appropriate material for use

as a lectotype was located, a neotype was selected. All

material cited has been seen unless it is expressly stated

otherwise. The JSTOR Plant Science website (http://

plants.jstor.org/) has been consulted in all applicable

cases and the Kew Herbarium Catalogue (http://apps.

kew.org/herbcat) was consulted for many species and

names as well. Data on localities is given as on the spec-

imens, with the present-day country where the specimen

was collected added.

RESULTS

The species are arranged alphabetically within the

four subgenera of Euphorbia that were established in

Bruyns et al. (2006). The synonymy is as in Bruyns et

al. (2006), except where otherwise mentioned and dis-

cussed.

NOMENCLATURAL ACCOUNT

1. Euphorbia subg. Chamaesyce Raf.

la. Sect. Anisophyllum Roeper

E. austro-occidentalis ThelL, Vierteljahrsschrift der

Naturforschenden Gesellschaft in Zurich 61; 431 (1916).

Type: Namibia, Okahandja, sandy bushveld, cultivated

land, 1 300 m, Oct., Dinter 105 (Z, lecto., designated

here; BOL, GRA, SAM-3 sheets, isolecto.). [Thellung

cited also Dinter 222 (Z), 222a (Z) and 822 (Z). A lecto-

type is selected.]

E. chamaesycoides B.Nord., Dinteria 11: 20 (1974).

Chamaesyce chamaesycoides (B.Nord.) Koutnik: 263

(1984). Type: Namibia, Brandberg, Upper Tsisab Valley,

± 1 600 m, 6 May 1963, Nordenstam 2567 (S, holo.; M,

iso.).

Bothalia 42,2 (2012)

Table 1. — Numbers of species, available (i.e. validly published) names and showing the numbers of species exhibiting different growth forms

(annuals, herbs, succulents, and geophytes) in the subgenera of Euphorbia.

E. eylesii Rendle, Journal of Botany 43; 52 (1905).

Chamaesyce eylesii (Rendle) Koutnik; 263 (1984).

Type: Zimbabwe, Deka siding along Bulawayo- Victoria

Falls railway line, May 1904, Evles 130 (BM, holo.;

SRGH, iso.).

E. leshumensis N.E.Br., Flora of Tropical Africa 6(1):

513 (1911). Type: Botswana, Leshumo Forest, received

May 1883, Holub (K, lecto., designated here). [Brown

(1911) also cited Macaulay 423 (K), from Zambia.]

E. glanduligera Pax, Botanische Jahrbiicher fur

Systematik 19: 142 (1894). Chamaesyce glanduligera

(Pax) Koutnik: 263 (1984). Type: Namibia, bei Nawas

am Swakop, bei Salem, 12 Dec. 1888, Giirich 3 (mis-

sing; sketch of type at K). Neotype (designated here):

Namibia, Naukluft Mtns, between Ababes and Flomnus,

Pearson 9106 (BOL).

E. pfeilii Pax.; 534 (1897). Type: Namibia, Stolzen-

fels, Rietfontein, 1890/1891, Pfeil 91 (missing).

E. glaucella Pax.: 737 (1898). Type: Namibia, Oka-

handja. Mar. 1883, Kopfner 68 (Z, lecto., designated

here). [Pax (1898) also cited Fleck 454a (Z), so a lecto-

type is selected.]

E. anomala Pax: 636 (1908), nom. illegit., non Bois-

sier ( 1862).

E. IvA’ebensis N.E.Br.: 137 (1909). Type; Botswana,

Kwebe Hills, 3300’, 7 Jan. 1897, Lugard 143 (K, lecto.,

designated here). [Brown (1909) also cited Lugard 81

(K), from the same locality and marked both specimens

as ‘Type’.]

E. gueinzii Boiss. in A.P. de Candolle, Prodromus

15(2): 71 (1862). Type: South Africa, at Natal Bay, Gue-

inzius (G, lecto., designated here; W, isolecto.). [Boissier

(1862) also cited an unnumbered collection of Sander-

son (in ‘h. Kew’, missing).]

E. gueinzii var. albovillosa (Pax) N.E.Br.; 252 (1915).

E. albovillosa Pax: 373 (1904). Type: South Africa,

Natal, Inchanga, 1 180 m, 16 Sept. 1893, Schlechter

3245 (BOL, lecto., designated here; GRA, K, PRE, iso-

lecto.). [A lectotype is designated as no specimens seen

by Pax have been located.]

E. inaequilatera Sond., Linnaea 23: 105 (1850). Ani-

sophyllum inaequilaterum (Sond.) Klotzsch & Garcke:

22 (I860). Chamaesyce inaequilatera (Sond.) Sojak:

169 (1972). Type: South Africa, Port Natal, Gueinzius

167 (MEL [501275], holo.; F, MEL, iso.).

Anisophyllum mundii Klotzsch & Garcke: 25 (1860).

Type: South Africa, Cape, Gamka R., Prince Albert div..

Jan. 1820, Mund & Maire 15 (K 000253186, lecto.,

designated here). [There are two specimens under this

number at K and it is not certain that either was seen

by Klotzsch & Garcke. This is the larger specimen, the

other is a ‘branch from the type’, according to N.E.

Brown.]

A. setigerum E.Mey. ex Klotzsch & Garcke: 29

(1860). Type: South Africa, Cape, Drege (missing).

E. parvifolia E.Mey. ex Boiss.: 34 (1862). Type:

South Africa, Cape, Jan. 1820, Mund & Maire 15 (K

000253186, lecto., designated here). [Boissier cited:

near Gariep, Drege; Beaufort distr., Lund; ‘Anis. Mun-

dtii Kl. et Gke l.c. p. 25’. From the latter, a lectotype is

selected.]

E. parvifolia var. laxa Boiss.: 34 (1862). Type: none

located. [Boissier cited: Drege 8191; 8198 and ‘Sieb.

Cap. n. 154, which have not been located.]

E. sanguinea var. setigera E.Mey. ex Boiss.; 35

(1862). Type; South Africa, near Kei and Bashee Rivers,

Drege (missing).

E. sanguinea var. natalensis Boiss.: 35 (1862). Type:

South Africa, Port Natal, Gueinzius 167 (F, lecto., des-

ignated here; MEL-2 sheets, isolecto.). [Boissier (1862)

did not say in which herbarium he had seen this speci-

men so a lectotype is selected.]

E. nelsii Pax: 737 (1898). Type: Namibia, Hereroland,

1886, L. Nels 91 (Z, holo.; K, iso.). [N.E. Brown anno-

tated the piece of Nels 91 at K as ‘fragment from type’

and mentioned also that the type was at Z.]

E. inaequilatera var. perennis N.E.Br.: 246 (1915).

Type: South Africa, Natal, near Tugela, 4 Jan. 1886,

Wood 3552 (K, lecto., designated here). [Brown also

cited many other syntypes for this variety.]

E. livida E.Mey. ex Boiss., in A.P. de Candolle, Pro-

dromus 15(2): 14 (1862). Chamaesyce livida (E.Mey. ex

Boiss.) Koutnik: 263 (1984). Neotype (designated here):

South Africa, Natal, without precise locality, Gerrard

1171 (K). [Boissier (1862) cited ‘ad Natal Bay, Drege'

and Gueinzius 177, which are both missing. The Drege

specimens found do not have this locality on them so a

neotype is selected.]

E. mossambicensis (Klotzsch & Garcke) Boiss. in

A.P. de Candolle, Prodromus 15(2): 36 (1862). Aniso-

phyllum mossambicense Klotzsch & Garcke: 30 (1860).

Chamaesyce mossambicensis (Klotzsch & Garcke)

Koutnik: 263 (1984). Type: Mo9ambique, Rios de Sena,

Peters 33 (K, lecto., designated here). [No type has been

Bothalia 42,2 (2012)

219

found at B. This is a fragment of the type so is desig-

nated as lectotype.]

E. neopolycnemoides Pax & K.Hoffm., Botanische

Jahrbiicher fur Systematik 45: 240 (1910). Chamae-

syce neopolycnemoides (Pax & K.Hoffm.) Koutnik: 263

(1984). Type: South Africa, Transvaal, between Nyl-

stroom and Naboomspruit, sandy places along Macha-

laquana R., 24 Jan. 1894, Schlechter 4278 (missing).

Neotype (designated here): South Africa, Waterberg,

Boschpoort near Warmbaths, 3650’, Jan 1906, Bolus

12280 (K; duplicate at BOL). [The type is missing and

the specimen at K of Bolus 12280 was matched by N.E.

Brown against Schlechter 4278 so it is selected as neo-

type.]

E. arabica van latiappendiculata Pax: 85 (1909).

Type: South Africa, Waterberg, Boschpoort near Warm-

baths, Jan 1906, Bolus 12280 (K, lecto., designated

here; BOL, isolecto.). [The specimen at K is ‘part of the

type’ and the other was not seen by Pax so a lectotype is

selected.]

E. pergracilis P.G.Mey., Mitteilungen aus der Bota-

nischen Staatssammlung Miinchen 6: 247 (1966).

Chamaesyce pergracilis (P.G. Mey.) Koutnik: 263

(1984). Type: Namibia, 7 miles east of Purros towards

Sesfontein, 23 June 1963, Giess 3211 (M, holo.; MO,

PRE, WAG, iso.).

E. phylloclada Boiss. in A.P. de Candolle, Prodromus

15(2): 66 (1862). Type: South Africa, between Verlept-

pram and mouth of Gariep, Sept., Drege 238 (S, lecto.,

designated here). [Boissier (1862) cited Drege ‘in h.

Bunge’ (missing), from the same locality.]

E. hereroensis Pax: 35 (1889). Type: Namibia, Here-

roland, Hykamkab, 300 m. May 1886, Marloth 1190

(missing).

E. rubriflora N.E.Br., Flora of Tropical Africa 6(1):

509 (1911). Type: Zimbabwe, Victoria Falls, Jan. 1906,

Allen 264 (K, lecto., designated here; SRGH, isolecto.).

[Brown (1911) also cited: Zambia, Livingstone, Rogers

7132 (K, BOL).]

E. schlechteri Pax, Botanische Jahrbiicher fiir Sys-

tematik 28: 26 (1900). Chamaesyce schlechteri (Pax)

Koutnik: 263 (1984). Type: Mozambique, Ressano

Garcia, 1 000’, 24 Dec. 1897, Schlechter 11915 (PRE,

lecto., designated here; BOL, BR, COI, G, GRA, HBG,

K, WAG, isolecto.). [No material definitely seen by Pax

in known and so a lectotype is selected.]

E. spissiflora S. Carter, Kew Bulletin. 45: 331 (1990).

Type: Zimbabwe, Nhongo, 8 km north of Gokwe, 6 Mar.

1964, Bingham 1158 (K, holo.; SRGH, iso.).

E. tettensis Klotzsch in W.C.H. Peters, Naturwis-

senschaftliche Reise nach Mossambique 1: 94 (1861).

Chamaesyce tettensis (Klotzsch) Koutnik: 263 (1984).

Type: Mozambique, Tete, Peters (missing). [The name

Anisophyllum tettense Klotzsch & Garcke: 34 (1860)

was not validly published since it only cited the above,

which had not yet appeared.]

E. zambesiana Benth., Hooker’s leones Plantarum

14: t. 1305 (1880). Chamaesyce zambesiaca (Benth.)

Koutnik: 263 (1984). Type: Malawi, Zomba Mtn, 1861,

Meller, Livingstone’s Expedition (K, lecto., desig-

nated here). [Bentham (1880) also cited: Malawi, Shire

Highlands, Blantyre, Buchanan 10 (K). A lectotype is

selected.]

lb. Sect. Articulofruticosae Bruyns

E. angrae N.E.Br., Flora capensis 5(2): 279 (1915).

Type: Namibia, Liideritz (Angra Pequena), 18 Jan. 1907,

Galpin & Pearson 7549 (K, lecto., designated here;

drawing; PRE, SAM, isolecto.). [Both the specimen at

SAM and that at K were annotated as ‘Type’ by N.E.

Brown, so a lectotype is selected.]

E. einensis G.Will.: 57 (2004). Type: Namibia, south-

ern Schakalberg, 70 km NE of Oranjemund, Williamson

5143 (BOL-2 sheets, holo.).

E. einensis var. anemoarenicola G.Will.: 62 (2004).

Type: South Africa, Kortdoom, Williamson 5985

(BOL-2 sheets, holo.).

E. burmannii E.Mey. ex Boiss. in A.P. de Candolle,

Prodromus 15(2): 75 (1862). Type: South Africa, Cape,

towards Blauwberg, Drege 2920 (P, lecto., designated

here). [Boissier (1862) cited Drege 2920 (P); Tygerberg,

Bergius; Krauss. Neither of the latter two specimens has

been located.]

E. biglandulosa Willd.: 27 (1814), nom. illegit. non

Desf. (1808). [Willdenow (1814) gave a description but

cited no specimens.]

Arthrothamnus burmannii E.Mey. ex Klotzsch &

Garcke: 62 (1860). Type: South Africa, Cape, Drege

(missing). Neotype (designated here): South Africa,

Cape, Drege 2920 (P). [No specimen as cited by

Klotzsch & Garcke (1860) has been found, so a neotype

is selected.]

Arthrothamnus bergii Klotzsch & Garcke: 63 (1860).

Type: South Africa, Cape, Bergius (missing).

E. phymatoclada Boiss.: 24 (1860). Type: South

Africa, rocky hills at Ebenezer, Drege 2943 (GRA,

lecto., designated here). [Boissier (1860) cited a speci-

men ‘in h. Bunge’, which has not been located so a lec-

totype with the same number is selected. This is from

a plant of E. burmannii, although this name is usually

placed as a synonym of E. mauritanica (e.g. White et al.

1941).]

E. hydnorae E.Mey. ex Boiss.: 95 (1862). Type:

South Africa, between Kaus and Doompoort, Drege

2943 (GRA, lecto., designated here). [Apart from

Drege 2943, Boissier (1862) also cited ‘in montibus

Niueweweld alt. 3 000^ 000 ped.’, apparently another

collection of Drege and both were ‘in h. Bunge’. The

lectotype selected here is a specimen of E. burmannii,

although this name is also usually placed as a synonym

of E. mauritanica (e.g. White et al. 1941 ).]

E. coiymbosa N.E.Br.: 279 (1915). Type: South

Africa, Cape, near Albertinia, 16 Nov. 1910, Muir (K,

holo.).

220

Bothalia 42,2 (2012)

E. kaiToensis (Boiss.) N.E.Br.: 290 (1915). E. bur-

mannii var. karroensis Boiss. in DC.: 75 (1862). Type:

South Africa, Cape, Karoo between Hoi River and

Mierenkasteel, 500-1 000’, 5 Aug. 1830, Drege 2947

(P, lecto., designated here; K, isolecto.). [Boissier

( 1 862) did not say in which herbarium the specimen was

located so a lectotype is selected.]

E. macella N.E.Br.: 288 (1915). Type: South Africa,

Cape, near Little Brak River, 10 Oct. 1814, Burchell

6197/2 (K, holo.).

E, ephedroides E.Mey. ex Boiss. in A.P. de Candolle,

Prodromus 15(2): 75 (1862). Type: South Africa, Cape,

Karoo at Goedemanskraal, 2 500’, 8 Sept. 1830, Drege

2949 (P, lecto., designated here; K, MO, S, isolecto.).

[Boissier (1862) cited also Burchell 1424 (‘in h. DC.’),

which has not been located.]

E. ephedroides var. immimita L.C. Leach & G.Will.:

72 (1990). Type: South Africa, Cape, Alexander Bay,

Williamson 3652 (NBG, holo.; K, PRE, iso.).

E. ephedroides var. debilis L.C. Leach; 73 (1990).

Type: Namibia, north of Rosh Pinah, Leach & Brimton

15893 (NBG, holo.; K, MO, PRE, iso.).

E. exilis L.C. Leach, South African Journal of Botany

56: 76 (1990). Type: South Africa, Cape, Aties, May

1984, Leach & Bayer 17129 (NBG-2 sheets, holo.; K,

iso.).

E. glandularis L.C.Leach & G.Will.: 75 (1990). Type:

South Africa, Cape, near Steinkopf, Leach & Hilton-

Tay lor 17019 (NBG, holo.; K, PRE, iso.).

E. gentilis N.E.Br., Flora capensis 5(2): 289 (1915).

Type; South Africa, Cape, Vanrhynsdorp Div., hills near

Zout River, 500’, 14 Jul. 1896, Schlechter 8136 (BOL,

lecto., designated here; GRA, HBG, K, PRE, S, iso-

lecto.). [Brown (1915) also cited Graafwater (Grauwa-

ter), 15 Dec. 1908, Pearson 3271 (BOL, K, SAM); near

Bitterfontein, Zeyher 1531 (G, K, S, W).]

E. vaalputsiana L.C.Leach: 534 (1988a). Type; South

Africa, Cape, Vaalputs, near Gamoep, Leach & Perry

1 7232 (NBG, holo.; K, MO, PRE, iso.).

E. gentilis subsp. tanquana L.C.Leach: 538 (1988a).

Type: South Africa, Cape, near turnoff to Skitterykloof,

Leach & Perry 17247a (NBG, holo.; K, M, MO, PRE,

iso.).

E. giessii L.C.Leach, Dinteria 16: 27 (1982). Type:

Namibia, 18 km east of Henties Bay, Dec. 1976, Giess

14809 {sub Leach 15940) (PRE, holo.; M, WIND, iso.).

E. herrei A. C. White et al.. The Succulent Euphor-

bieae 2: 962 (1941). Type: South Africa, Cape, near

Swartwater, 1930, llerre sub PRE 46025 (PRE, holo.).

[Although White et al. (1941) did not mention the

number PRE 46025, it is assumed that this is the same

specimen as the one they cited.]

E. juttae Dinter, Neue und wenig bekannte Pflanzen

Deutsch-SWA’s: 30 (1914). Type: Namibia, Garub, 900

m, 9 Jan. 1910, J. Dinter 1047 (SAM, lecto., designated

by Leach (1988a); NY, isolecto.).

E. siliciicola Dinter: 31 (1914). Type: Namibia, Bull-

sport, 5 Apr. 1911, Dinter 2132 (SAM, lecto., designated

by Leach (1988a)).

E. aequoris N.E.Br.: 279 (1915). Type: South Africa,

Cape, Middelburg div., Schoombie, Feb. 1897, Trol-

lip (sub SAM 20091) (SAM, lecto., designated here; K,

isolecto.). [Brown (1915) cited also: Rosmead Junc-

tion, 4 000’, 22 Mar. 1900, Sim sub Galpin 5626 (PRE);

between Colesburg & Hanover, 1871, Bolus 2201 (K).]

Leach (1988a) discussed E. juttae in detail but con-

sidered that E. aequoris, although closely related, was

‘suffieiently distinct in vegetative characters and habit

alone for it to be disregarded’ in those discussions. He

did not say in what way it was so distinct. It appears

that this distinctiveness lay in the much more robust

plants formed by E. aequoris, with longer and more

slender stems and branches, with more widely spaced

and less prominent tubercles and a longer rootstock, as

well as the lack of the peculiar habit that the branches

have in E. juttae of bending over to the north or west.

Nevertheless, among the material that he cited under E.

juttae were two specimens from near Olifantshoek and

near Kenhardt respectively that are rather more typical

of E. aequoris than of E. juttae. While many specimens

of E. aequoris are unmistakeable (especially those from

the Great Karoo and drier parts of the Eastern Cape),

those from the Northern Cape and calcareous pans on

the southern edge of the Kalahari are not clearly refer-

able to either species. Some of these (especially plants

from exposed spots) may even exhibit a similar, almost

prostrate habit to E. juttae, and have shorter stems and

branches with more prominent tubercles while more pro-

tected plants are erect, slender, and more typical of E.

aequoris. I have found no clear distinctions between the

two species and have placed E. aequoris in synonymy.

E. lavrani L.C.Leach, The Journal of South African

Botany 49: 807 (1983). Type: Namibia, Namuskluft,

1 200 m, Lavranos & Newton 16872 (PRE holo.; NBG,

SRGH, iso.).

E. muricata Thunb., Prodromus plantarum capen-

sium 2: 86 (1800). Type: South Africa, Cape, Thunberg

(UPS-THUNB 11499, holo.; drawing and fragment at K,

iso.).

E. spicata E.Mey. ex Boiss.: 97 (1862). Type: South

Africa, 31 Aug. 1830, Drege 2946 (K, lecto., designated

here; S, isolecto.). [Boissier (1862) cited also Cape, near

Bitterfontein, Zeyher 1531 (G, K, S, W), which is E.

gentilis.)

E. aspericaulis Pax: 26 (1899). Type: South Africa,

Cape, Hantam Mtns, ex Dr Meyer (holo. missing; draw-

ing and fragment at K, iso.). [According to Carter (2002)

this specimen is at B, but it cannot be located.]

E. rhombifolia Boiss., Centuria Euphorbiarum: 19

(1860). Type: South Africa, Cape, arid places on south-

ern Karoo, Drege 8217 (G, lecto., designated here; K, S,

W, isolecto.). [Boissier (1860) cited also Ecklon & Zey-

her-. Euphorb. 23, 83 (G, W).]

Arihr-othamnus densiflor-us Klotzsch & Garcke: 62

(1860). Type: South Africa, Cape, Karoo near Olifants

Bothalia42,2 (2012)

221

River, Oudtshoom distr., Jan. 1820, Mimd & Maire (K,

lecto., designated here). [Since there is no evidence that

Klotzsch & Garcke saw this specimen it is designated as

lectotype.]

E. brachiata (E.Mey. ex Klotzsch & Garcke)

Boiss.: 74 (1862). Arthrothamnus brachiatus E.Mey.

ex Klotzsch & Garcke: 62 (1860). Type: South Africa,

Cape, near Ebenezer, Drege 2948 (K, lecto., designated

here; S, isolecto.). [Although Boissier (1862) included

E. miiricata Thunb. in the synonymy of E. brachiata,

this is considered here as a separate species. Since it is

unlikely that Klotzsch & Garcke saw any of the sheets

listed, a lectotype is selected. The specimen at K of

Drege 2948 was taken from ‘the type’ in ‘Drege’s her-

barium’ by N.E. Brown.]

E. decussata E.Mey. ex Boiss.: 74 (1862), nom ille-

git., non Salisb. (1796). [Boissier (1862) cited Drege

3926 (missing), Drege 8218 (K, MO, S, W) and ‘Oli-

fants River, Mund & Maire' (K). However, since it is an

illegitimate name, a lectotype is not selected here.]

E. amarifontana N.E.Br.: 275 (1915). Type: South

Africa, Cape, near Springbokkuil River, Bitterfontein,

Zeyher 1534 (K, lecto., designated here; BOL, SAM

‘1534b’-2 sheets, isolecto.). [Brown (1915) cited also

Pearson 5532 (BOL).]

E. chersina N.E.Br.: 274 (1915). Type: Namibia,

Liideritz (Angra Pequena), 18 Jan. 1907, Galpin &

Pearson 7584 (K, lecto., designated here; PRE, iso-

lecto.). [Brown (1915) also cited Marloth 4638 (K)

and marked both specimens as ‘Type’, so a lectotype is

selected.]

E. caterviflora N.E.Br.: 286 (1915). Type: South

Africa, Cape, Nieweveld, Beaufort West, Drege 8218

(K, lecto., designated here; G, MO, W, isolecto.).

[Brown (1915) also cited and wrote ‘Type’ on Tyson 167

(K, SAM), so a lectotype is selected.]

E. hastisquama N.E.Br.: 288 (1915). Type: South

Africa, Cape, fields by the Swartkops River, Zey-

her 1099 (BOL, lecto., designated here; K, isolecto.).

[Brown (1915) cited also Zeyher 3854 (SAM) and Eck-

lon & Zeyher, Enphorb. 25 (K, SAM).]

E. mundii N.E.Br.: 287 (1915). Type: South Africa,

Cape, Montagu, 1 Jan. 1903, Marloth 2805 (K, lecto.,

designated here; PRE, isolecto.). [Brown (1915) also

cited Marloth 3904 (PRE) and Marloth 4878 (K)

and several others as syntypes. Brown gave this as a

new name for Arthrothamnus densiflorus Klotzsch &

Garcke, which could not be transferred to Euphorbia.

He regarded these syntypes as identical to the Mund &

Maire specimen that typified A. densiflorus and therefore

named this plant after Mund.]

E. perpera N.E.Br.: 277 (1915). Type: South Africa,

Cape, along Orange River, between Verleptpram and its

mouth, Drege (K, holo.).

E. rudolfii N.E.Br.: 276 (1915). Type: South Africa,

Cape, Vanrhynsdorp div., Bitterfontein, Sept. 1897,

Schlechter 11047 (K 000252612, lecto., designated here;

BR, GRA, K. L-2 sheets, PRE, S, isolecto.). [Brown

cited also: between Bitterfontein and Stinkfontein, 5

Dec. 1910, Pearson 5533 (BOL, K). He only wrote

‘Type’ on the specimens at K of Pearson 5533 and

on one of the specimens at K of Schlechter 11047 (K

000252612). Therefore a lectotype is selected here.]

E. bayeri L.C. Leach: 539 (1988b). Type: South

Africa, Cape, 2 km west of Mossel Bay, 11 Sept. 1985,

Bayer 4875 (NBG, holo.; K, MO, PRE, iso.).

E. spartaria N.E.Br. Flora of Tropical Africa 6(1):

558 (1911). Type: Namibia, Hoffnung, Feb. 1907,

Galpin & Pearson 7560 (K, holo.; PRE, SAM, iso.).

[Brown annotated the sheet of Galpin & Pearson 7560

(K) as ‘Type’ and that at SAM as ‘Part of the type’. He

did not do this for any of the specimens of Dinter 983

(K, SAM-2 sheets) designated by Leach & Williamson

(1990) as lectotype and also not on Dinter 255 (K). Con-

sequently their lectotype is set aside here in favour of

Brown’s preferred ‘type’.]

E. racemosa E.Mey. ex Boiss.: 75 (1862), nom. ille-

git., non Tausch ex Rchb. (1832). [Boissier (1862)

cited: South Africa, Cape, near Hamerkuil, Drege (MO,

S ‘8204’); distr. Beaufort, Ecklon in h. Petrop (miss-

ing). As this is an illegitimate name a lectotype is not

selected.]

E. indecora N.E.Br.: 274 (1915). Type: South Africa,

Cape, between Dabenoris and Houms Drift, 1 1 Jan.

1909, Pearson 3387 (K 000252597, lecto., designated

here; K, isolecto.). [There are two specimens of this col-

lection at K, both annotated by N.E. Brown as ‘Type’,

so a lectotype is selected. The size of these plants (2-3’

tall, according to the specimens) suggests that they are

E. spartaria rather than E. rhombifolia.]

E. rhombifolia var. laxa N.E.Br.: 285 (1915). Type:

South Africa, Cape, among rocks along Chichaba River

between Komgha and Kei Mouth, Aug. 1891, 1 000’,

Flanagan 838 (GRA, lecto., designated here; PRE,

SAM, isolecto.). [In this case (unlike for E. spartaria)

Brown annotated the sheets of the different collections

MacOwan 1612 (GRA) and Flanagan 838 (GRA, PRE,

SAM) as ‘type’ so a lectotype is designated. Another

syntype is Sutherland (K).]

E. rhombifolia var. triceps N.E.Br.: 285 (1915). Type:

South Africa, Cape, Queenstown distr., mountains near

Imbasa River, 1860, Cooper 318 (K, lecto., designated

here; BOL, W, isolecto.). [Brown (1915) cited several

specimens as representing var. triceps, so a lectotype is

selected.]

E. cibdela N.E.Br.: 275 (1915). Type: Namibia, on

hills at Schakalskuppe, 4 900-5 600’, 18 Jan. 1909,

Pearson 4428 (K, holo.; BOL, ED, SAM, iso.).

E. recfirawa N.E.Br.: 283 (1915). Type: South Africa,

Cape, Klipfontein, Griqualand West, 29 Dec. 1812,

Burchell 2633 (K, lecto., designated here). [Brown

(1915) cited several specimens, of which the above is

selected as lectotype.]

E. spinea N.E.Br., Flora capensis 5(2): 272 (1915).

Type: Namibia, among rocks near Dabegabis, Pear-

son 4380 (K, lecto., designated here; BOL, isolecto.).

222

Bothalia 42,2 (2012)

[Brown (1915) also cited Pearson 3296 (BOL, K,

SAM), which is E. rhombifolia and Pearson 4585 (K)

and wrote ‘Type’ on all three specimens, so a lectotype

is designated.]

E. stapelioides Boiss., Centuria Euphorbiarum; 26

(1860). Type: South Africa, Cape, at the mouth of the

Gariep (Orange), 4 Oct. 1830, Drege 8199 (P, holo.; S,

W, iso.).

E. lumbricalis L.C. Leach: 369 (1986b). Type: South

Africa, Cape, north of Koekenaap, 10 May 1984, Leach

& Bayer 17123 (NBG, holo.; K, MO, PRE-2 sheets,

iso.).

E. suffulta Bruyns, South African Journal of Botany

56: 129 (1990). Type: South Africa, Cape, Tierberg,

Prince Albert distr., 6 Dec. 1987, Bruyns 2902 (BOL,

holo.; K, PRE, iso.).

E. tenax Burch. Travels in the Interior of southern

Africa, 1: 219 (1822). Type: South Africa, Cape, Hang-

klip, near Ongeluks River, Ceres div., 17 July 1811,

Burchell 1219 (K, holo.).

E. arceuthobioides Boiss.: 20 (1860). Type: South

Africa, Cape, 70.10, Ecklon & Zeyher, Euphorb. 76,

{Ecklon 1312) (G, holo.; W, iso.).

Arthrothamnus ecklonii Klotzsch & Garcke: 63

(1860). Type: South Africa, Cape, Ecklon & Zeyher,

Euphorb. 24, {Ecklon 1871) (W, lecto., designated here).

[Klotzsch & Garcke (1860) cited ‘Ecklon n. 23. 25 & 24

ex parte’. The only specimens from the Ecklon and Zey-

her collections with similar numbering are those labelled

‘Euphorb. 23’, ‘Euphorb. 24’ and ‘Euphorb. 25’ and so it

must be to these that Klotzsch & Garcke referred.]

Arthrothamnus scopiformis Klotzsch & Garcke: 63

( 1 860). Type: South Africa, Cape, Bergius (missing).

E. rhombifolia var. cymosa (Klotzsch & Garcke)

N.E.Br.: 285 (1915). Arthrothamnus cymosus Klotzsch

& Garcke: 63 (1860). Type: South Africa, Cape, Eck-

lon & Zeyher, Euphorb. 24 (W, lecto., designated here).

[Klotzsch & Garcke (1860) cited ‘Ecklon n. 24 ex

parte’. This is assumed to be the same as "Ecklon & Zey-

her, Euphorb. 24' , of which there is a piece in W. This

piece belongs to E. tenax. However, there is no evidence

that they saw this specimen and so it is selected as a lec-

totype.]

E. serpiformis Boiss.: 75 (1862). Type: South Africa,

Cape, Berg River Valley, Zeyher 1535 (BOL, lecto., des-

ignated here; K, S, SAM, W, WU, Z, isolecto.). [Bois-

sier (1862) cited also ‘Eckl. & Zeyh. 24’ (i.e. Ecklon &

Zeyher, Euphorb. 24 (W)) and ‘Riesvallei {Bergius h.

Berol.y (missing).]

E. mixta N.E.Br.: 585 (1925). E. arrecta N.E.Br.: 283

(1915), nom. illegit., non N.E.Br. (1914). Type: South

Africa, Cape, Berg River Valley, Zeyher 1535 (K, holo.;

BOL, S, SAM, W, WU, Z, iso.).

In Bruyns et al. (2006), E. tenax was treated as a syn-

onym of E. arceuthobioides. The respective types make

it clear that they are the same species. However, E. tenax

was published first and so this treatment was wrong.

E. verruculosa N.E.Br., Flora capensis 5(2): 585

(1925). Type: Namibia, Luderitz (Angra Pequena), 10

miles from coast, Nov. 1908, Marloth 4639 (PRE, holo.;

K, iso.). [Brown annotated the specimen at PRE as

‘Type’ and that at K as ‘half of the Type sheet, presented

to Kew by Dr Marloth’. So the sheet at PRE is taken as

the holotype.]

lc. Sect. Espinosae Pax & K.Hoffm.

E. guerichiana Pax, Botanische Jahrbucher fur Sys-

tematik 19: 143 (1894). Type: Namibia, rocks south of

Khorixas, 14 Nov. 1888, Gurich 73 (missing). Neotype

(designated here): Namibia, Ababes, banks of Tsondap

River, 30 Dec. 1915, Pearson 9119 (BOL).

E. commiphoroides Dinter: 90 (1909). Neotype (des-

ignated here): Namibia, Tsumeb distr., Auros, 10 Feb.

1925, Dinter 5596 (BOL; duplicate at SAM). [Dinter

(1909) cited no specimens and only mentioned ‘Haufig

in Hererolande: Salem, Modderfontein, Omburo, Tsao-

bis, Omatako’. No specimens from any of these locali-

ties have been found. A neotype is therefore selected.]

E. frutescens N.E.Br.: 270 (1915). Type: Namibia,

lower mountain slopes of Aus, 3 000’, Jan. 1909, Pear-

son 4714 (K, holo.; BOL, SAM, iso.). [Although several

of these sheets are labelled ‘Type’, only that at K was

annotated by N.E. Brown himself and so this specimen

is taken as the holotype.]

E. espinosa Pax, Botanische Jahrbucher fur Sys-

tematik 19: 120 (1894). Type: Tanzania, without precise

locality, Fischer 285 (K, lecto., designated here). [No

material definitely seen by Pax in known and so a lecto-

type is selected.]

E. gynophora Pax: 374 (1904). Type: Tanzania, Pare

Mountains, befweem Kisuani and Madji-ya-juu, 700 m,

13 Oct. 1902, Engler 1579 (K, drawing, lecto., desig-

nated here). [Pax (1904b) cited Engler 1579 and Engler

1586. No material definitely seen by Pax is known, but

drawings of both these specimens are at K. One of these

is selected as lectotype.]

ld. Sect. Erondosae Bruyns

E. leistneri R.Archer, South African Journal of Bot-

any 64: 258 (1998). Type: Namibia, east of Epupa Falls,

Jul. 1976, Leistner et al. 264 (PRE, holo.; B, K, WIND,

iso.).

E. transvaalensis Schltr., Journal of Botany 34: 394

(1896). Type: South Africa, Transvaal, near Edwin Bray

Battery, shady kloofs in Kap River Valley, Barberton,

2 000’, fl. Nov. 1890, Galpin 1198 (GRA, lecto., desig-

nated here; K, NH, SAM, Z, isolecto.). [Since there is no

sign that Schlechter saw any of the sheets listed, a lec-

totype is selected. Brown compared the specimen at K

with the type, but did not state where the latter was.]

E. galpinii Pax: 742 (1898). Type: South Africa,

Transvaal, near Edwin Bray Battery, Barberton, 2 000’,

fl. Nov. 1890, Galpin 1198 (SAM, lecto., designated

here; GRA, K, NH, Z, isolecto.). [No material definitely

seen by Pax in known and so a lectotype is selected.]

Bothalia 42,2 (2012)

223

E. ciliolata Pax: 743 (1898). Type: Angola, Sierra

Chella and Gambos, 900-1 100 m, Antimes & Dekindt

781 (BR, lecto., designated here; LISC, Z, isolecto.).

[No material definitely seen by Pax in known and so

a lectotype is seleeted. Specimens labelled "Dekindt

757 ’are at Z and BR while at LISC there is a specimen

labelled "Antimes 78 E. These are all assumed to be the

same collection, namely Antimes & Dekindt 757.]

2. Euphorbia subg. Esula Pers.

E. albanica N.E.Br., Flora capensis 5(2): 258 (1915).

Type: South Africa, Albany div., Brookhuisens Poort,

near Grahamstown, MacOwan 657 (GRA, holo.; K, iso.)

E. berotica N.E.Br., Flora of Tropical Africa 6(1):

600 (1912). Type: Angola, Mo9amedes distr., foot of

Sierra Negros, behind the mouth of the Bero River, July

1859, Welwitsch 633 (BM, holo.; LISU, iso.).

E. epicyparissias E.Mey. ex Boiss. in A.P. de Can-

dolle, Prodromus 15(2): 168 (1862). Type: South Africa,

Transvaal, near Vaal River, Burke (K, lecto., designated

here). [Boissier (1862) cited: Cape, near Zwangerberg,

Drege; Mimd & Maire in h. Berol; near Vaal R, in h.

Kew, Burke (K). The lattermost is selected as lectotype.]

Tithymalus epicyparissias E.Mey. ex Klotzsch &

Garcke: 88 (1860). Type: South Africa, Cape, Drege

(HBG, holo.; MO, W-3 sheets, iso.).

Tithymalus involiicratus E.Mey. ex Klotzsch &

Garcke: 91 (1860). Type: South Africa, Drege (HBG,

lecto., designated here; MO, isolecto.). [Klotzsch &

Garcke ( 1 860) cited: Drege (MO, HBG), Ecklon & Zey-

her n. 6; Ecklon & Zevher n. 8 (HBG, S, SAM); Krebs

(K).]

E. involucrata E.Mey. ex Boiss.: 168 (1862). Type:

South Africa, near Phillipstown, Ecklon & Zeyher n.

8 (HBG, lecto., designated here; S, SAM, isolecto.).

[Boissier (1862) cited: near George, Drege (BM, MO);

between Langekloof and ‘Zoega’ R., Krauss; near Phil-

lipstown, Ecklon & Zeyher n. 6; Ecklon & Zeyher n. 8

(HBG, S, SAM).]

E. bachmannii: Pax: 535 (1897). Type: South Africa,

Pondoland, end Oct. 1888, Bachrnann 755 (missing).

E. involucrata var megastegia Boiss.: 168 (1862).

Type: South Africa, Cape, near Katberg, Drege, Krebs.

E. epicyparissias var. puberula N.E.Br.: 267 (1915).

Type: South Africa, Kentani, 1 200’, 8 Oct. 1910, Pegler

460 (K, holo.; SAM, iso.).

E. epicyparissias var. wahlbergii (Boiss.) N.E.Br.:

267 (1915). E. wahlbergii Boiss.: 169 (1862). Type:

South Africa, 1842, Wahlberg (S, lecto., designated

here). [Boissier (1862) cited: ‘South Africa, between

Umtata and Omgaziana, Drege, Wahlberg, h. Bunge &

Holm’. The latter is in S.]

E. ericoides Lam., Encyclopedic methodique 2(2):

430 (1788). Type: South Africa, Cape of Good Hope,

Sonnerat (P-LAM P00381881, holo.).

E. erythrina Link, Enumeratio plantarum horti regii

berolinensis altera 2: 12 (1822). Tithymalus erythri-

nus (Link) Klotzsch & Garcke: 91 (1860). Type: South

Africa, Cape of Good Hope, Bergius (missing). Neotype

(designated here): South Africa. Cape, Paarl Mountain,

Drege 2197 (K 000253220; duplicate at K).

E. erythrina var. meyeri N.E.Br.: 262 (1915). E. mey-

eri Boiss.: 35 (1860), nom. illegit., non Steud. (1840).

Type: South Africa. Cape, Paarl Mountain, Drege 2197

(K 000253220, lecto., designated here; K, isolecto.).

[Since E. meyeri Boiss. was illegitimate, I treat var. mey-

eri as described by Brown. Brown (1915) cited several

specimens: without locality, Mimd & Maire, Malmes-

bury, Schlechter 5348; Paarl Mountain, Drege 2197 (K);

mountains near Cape Town, Ecklon & Zeyher Euphorb.

14 (LE).]

Tithymalus apiculatus Klotzsch & Garcke: 94 (1860).

Type: South Africa. Cape, Mimd & Maire (K, lecto.,

designated here). [Cited were: South Africa. Cape, Eck-

lon & Zeyher 14 (LE); Mimd & Maire (K) so a lecto-

type is selected. The latter is annotated by N.E. Brown

as ‘from the type’ from the Berlin Herbarium.]

Tithymalus confertus Klotzsch & Garcke: 94 (1860).

Type: South Africa. Cape, Mimd & Maire (K, lecto.,

designated here). [Cited were: South Africa. Cape,

Ecklon & Zeyher 5 (SAM); Mimd & Maire (K) so a

lectotype is selected. The specimen Ecklon & Zey-

her 5 (SAM) is of E. ericoides rather than E. ery-

thrina (though the label on it gives "Euphorbia striata

Thunb.’).]

E. erythrina var. burchellii Boiss.: 169 (1862). Type:

South Africa, Burchell 458 (missing). [This specimen

was said to be ‘in herb. DC.’]

E. foliosa (Klotzsch & Garcke) N.E.Br. Flora ca-

pensis 5(2): 262 (1915). Tithymalus foliosus Klotzsch

& Garcke: 67 (1860). Type: South Africa, Cape Flats,

near Cape Town, Ecklon & Zeyher 12 (K 000253222,

lecto., designated here; K, SAM, isolecto.). [The type of

Klotzsch & Garcke has not been located but Brown kept

part of it at K (comment on 000253222).]

E. dumosa E.Mey. ex Boiss.: 168 (1862), nom. ille-

git., non A. Rich. (1850). Types: South Africa, Pondo-

land, near the Umsikaba River, Drege 4619 (K, 2 sheets,

MO); "Eckl. & Zeyh 86' (missing). [Since this is an ille-

gitimate name, a lectotype is not selected here.]

E. artifolia N.E.Br.: 263 (1915). Type: South Africa,

Milkwoodfontein, Riversdale div., ± 600’, 7 Oct. 1897,

Galpin 4562 (K, holo.; PRE, iso.). [The specimen at K

was annotated as ‘Type Specimen’ by N.E. Brown while

that at PRE was not annotated by him. Consequently the

one at K is the holotype.]

E. genistoides PJ. Bergius, Descriptiones Plantarum

ex Capite Bonae Spei: 146 (1767). Tithymalus gen-

istoides (P.J.Bergius) Klotzsch & Garcke: 97 (1860).

Galarhoeus genistoides (P.J.Bergius) Haw.: 144 (1812).

Type: South Africa, Cape of Good Hope, Auge (Grubb)

(SBT 3.1.6.13, holo.).

Bergius only cited ‘Herm. Aff. 23’, which refers to

page 23 in J. Burman’s Catalogi duo plantarum afri-

canorum of 1736 that was in turn part of his Thesaurus

224

zeylanicus. No illustration or specimen is listed, only a

phrase which corresponds to the same phrase on page

23 in Burman’s Catalogi. However, there is a specimen

at SBT annotated by Bergius as ‘Euphorbia mihi gen-

istoides’ and ‘e. Cap. b. sp. Grubb’. It is known that a

consignment of specimens collected at the Cape by J.A.

Auge was bought from Auge by Michael Grubb during a

brief visit to the Cape in 1764 and presented to Bergius,

and that these formed the basis of Bergius’ "Descrip-

tiones’ (Gunn & Codd 1981). Consequently, this speci-

men is taken as the type. Haworth (1812) did not refer

to Bergius’ publication directly, but to ‘Willd., Sp. PI. 2:

908’ where references ‘Mant. 564’ and ‘Berg. cap. 146’

were given, the latter clearly the same as above.

Tilhymahis revolutus Klotzsch & Garcke: 99 (1860).

Type: South Africa. Cape of Good Hope, Ecklon & Zey-

her 2 (missing).

E. genistoides var. puberiila N.E.Br.: 264 (1915).

Type: South Africa, Cape, Lion Mountain, Wolley-

Dod 3104 (K, lecto., designated here; BOL, isolecto.).

[Brown (1915) cited: without locality, Thimberg; Mimd;

Harvey 444 (K); near Hopefield, Bachmann 55; New

Kloof, Drege\ Lion Mountain, Drege 8192 (HBG); Sch-

lechter 1381; Wolley-Dod 3104 (BOL, K ); near Cape

Town, Prior (K); Simon’s Bay, Wright 447.1

E. genistoides var. corifoUa (Lam.) N.E.Br.: 264

(1915). E. corifolia Lam.: 431 (1788). Type: South

Africa, Cape of Good Hope, Sonnerat (P-LAM

P00381882, holo.;K, iso.).

E. kraussiana Bernh. ex C.Krauss, Flora 28: 87

(1845). Type: South Africa, Natal, forest margins near

Pietermaritzburg, Sept. 1839, 2 000-2 500’, Krauss 256

(MO, holo.; BM, K-2 sheets, iso.). [Bemhardi’s herbar-

ium was bought by MO (Gunn & Codd 1981) and, since

Bemhardi drew up the description and Krauss published

it, the holotype is taken as the specimen at MO.]

Tithymalus truncatus Klotzsch & Garcke: 75 (1860).

Type: South Africa, Cape, Krebs (missing).

Tithymalus meyeri Klotzsch & Garcke: 75 (1860).

Type: South Africa, Cape, Ecklon & Zeyher Euphorh. 13

(Z, lecto., designated here; SAM, isolecto.). [Klotzsch

& Garcke (1860) also cited 'Drege’ and 'Krebs’, which

have not been located.]

E. kraussiana var. erubescens (E.Mey. ex Boiss.)

N.E.Br.: 268 (1915). E. erubescens E.Mey. ex Boiss.:

116 (1862). Type: South Africa, Natal, between Um-

zimkulu & Umkomaas, Apr., Drege (S, lecto., des-

ignated here; BM, isolecto.). [Boissier (1862) cited

‘Zuurbergen (‘2347’ K); near Grahamstown, Drege (K);

near Vanstadensriver, Krauss; between Umzimkulu &

Umkomaas, Drege (BM, S); ‘Winterberg, Ecklon & Zey-

her'. Only that at S is annotated by Boissier.]

E. mauritanica L., Species Plantarum 1: 452 (1753).

Tithymalus mauritanicus (L.) Haw.: 139 (1812). Type:

Illustration in Dillen., Hort. Eltham. 2: 384, t. 289, f 373

( 1 732) (lecto., designated by Croizat 1945).

Tithymalus zeyheri Klotzsch & Garcke: 71 (1860).

Type: South Africa, Cape, Ecklon & Zeyher, Euphorh.

26 (missing).

Bothalia 42,2 (2012)

T. brachypus Klotzsch & Garcke: 74 (1860). Type:

South Africa, Cape, Bergius (missing).

E. melanosticta E.Mey. ex Boiss.: 95 (1862). Type:

South Africa, Kaus Mountain, towards Goedemanskraal,

2 500’, Drege 2945 (K, lecto., designated here; MO, iso-

lecto.). [Boissier (1862) cited a specimen at ‘h. Bunge’

that has not been located, so a lectotype is selected.]

E. mauritanica var. namaquensis N.E.Br.: 292 (1915).

Type: South Africa, Pofadder distr., Groot Rosynbos,

9 Jan. 1909, Pearson 3845 (K, lecto., designated here;

BOL, NBG, Z, isolecto.). [Brown (1915) cited (among

others): Namibia, koppie about 20 km south of Warm-

bad, 27 Jan. 1909, Pearson 4432 (BOL, K); South

Africa, between Groot Rosynbos and Wortel, 10 Jan.

1909, Pearson 3628 (BOL, K).]

E. sarcostemmatoides Dinter: 304 (1921b). Type:

Namibia, (Tsamkubis ?) Klein Aub, 7 Apr. 1911, Dinter

2149 (SAM, lecto., designated here). [Dinter (1921b)

cited 2 collections: Dinter 2149 (SAM) and 2532a

(missing).]

E. paxiana Dinter: 265 (1921a). Type: Namibia, Klein

Aub, am schwarzem Kam Rivier im Bastardland, Dinter

2652 (SAM, holo.).

E. mauritanica var. foetens Dinter ex A. C. White et

al.\ 961 (1941). Type: Namibia, 8 km east of Pomona,

14 June 1929, Dinter 6418 (PRE, holo.; BOL, HBG-2

sheets, K, M, NBG, S, SAM, iso.).

E. mauritanica var. minor A.C. White et al: 961

(1941). Type: South Africa, Cape, 30 miles north of

Laingsburg, Aug. 1939, Dyer 4105 (PRE, holo.; K, iso.).

E. mauritanica var. lignosa A.C. White et al.: 961

(1941). Type: Namibia, Namib near Liideritzbucht, Nov.

1908, Marloth 4638 (PRE 0248633-0, holo.; PRE, iso.).

E. mauritanica var. corallothamnus Dinter ex

A.C. White et al: 961 (1941). Type: Namibia, dunes

near Buchuberge, 1 July 1929, Dinter 6467 (PRE, holo.;

BOL, HBG-3 sheets, K, LD, M, NBG, S, SAM, iso.).

E. muraltioides N.E.Br., Flora capensis 5(2): 264

(1915). Type: South Africa, Albany div., Brookhuisens

Valley, MacOwan 642 (K, lecto., designated here; GRA,

isolecto.). [Brown (1915) also cited MacOwan 329

(GRA, K) and Glass 665 (K, SAM) and wrote ‘Type’ on

all of them.]

E. natalensis Bernh. ex Krauss Beitrage zur Flora des

Cap- und Natallandes: 150 (1846). Type: South Africa,

Natal, base of Tafelberg, Aug. 1839, Krauss 434 (MO,

holo.; BM, FI, K, M, iso.). [Krauss (1845) mentioned

the number '434' , though this did not appear in Krauss

( 1 846). As for E. kraussiana, the holotype is at MO.]

Tithymalus capensis Klotzsch & Garcke: 98 (1860).

Type: South Africa, Cape of Good Hope, Ecklon & Zey-

her (missing), Drege (missing).

E. ruscifolia (Boiss.) N.E.Br., Flora capensis 5(2):

259 (1915). E. sclerophylla var. ruscifolia Boiss.: 169

(1862). Type: South Africa, between Kei and Gekau,

Drege 4621 (missing). Neotype (designated here): South

Bothalia 42,2 (2012)

225

Africa, Cape, Krielis Country, Bowker (K). [Boissier

(1862) cited a specimen at ‘h. Bunge’ that has not been

located, so a neotype is selected. This was compared by

N.E. Brown with Drege 4621 in Liibeck.]

E. sclerophylla Boiss., Centuria Euphorbiarum: 37

(1860). Type: South Africa, Cape, ad Grahamstown, Jul.

1829, Ecklon et Zeyher n° II (G, lecto., designated here;

LE, MO (only piece on right hand side), SAM, W, iso-

lecto.). [Boissier (1860) cited: Ad. Prom. B. spei, Krebs

pi. exs. n° 296 (G-DC, LE); ad Grahamstown, Ecklon &

Zeyher n° 11 (G, LE, SAM, W).]

Tithymahis multicaiilis Klotzsch & Garcke: 98

(1860). Type: South Africa, Cape of Good Hope, Krebs

(missing).

E. ovata (E.Mey. ex Klotzsch & Garcke) Boiss.:

167 (1862). Tithymalus ovatus E.Mey. ex Klotzsch &

Garcke: 97 (I860). Type: South Africa, Cape of Good

Hope, Drege (ED, lecto., designated here; MO, NY, iso-

lecto.). [A lectotype is designated as it cannot be ascer-

tained whether Klotzsch & Garcke saw any of these

sheets.]

E. sclerophylla var. myrtifolia E.Mey. ex Boiss.: 169

(1862). Type: South Africa, near Assegaaibosch, Drege

3563 (P, holo.; K-2 sheets, iso.). [Sheets at HBG and

MO do not have the number ‘35(53’ on them and are not

included here.]

E. striata var. brachyphylla Boiss.: 170 (1862). Type:

South Africa, Sterkstroom div., plains on top of Katberg,

Drege (K 000253210, lecto., designated here). [Boissier

(1862) cited ‘South Africa, Sterkstroom div., plains on

top of Katberg, Drege (K 000253210); Los Tafelberg,

5 000-6 000’, Drege’ (missing). The first specimen is of

E. ovata = E. sclerophylla and so selecting it as the type

means that this name becomes a synonym of E. sclero-

phylla rather than of E. striata.]

E. sclerophylla var. piiberiila N.E.Br.: 260 (1915).

Type: South Africa, Bathurst div., Rietfontein, between

Kariega River and Port Alfred, Biirchell 3961 (K, holo.).

E. stolonifera Marloth ex A. C. White et ah. The Suc-

culent Euphorbieae 2: 961 (1941). Type: South Africa,

Cape, near Matjiesfontein and ‘Dwars in die Weg’, 900

m, Oct. 1920, Marloth 9836 (PRE 0838532-0, holo.;

PRE, iso.).

E. striata Thimb., Prodromus plantarum capensium

2: 86 (1800). Tithvmalus striatiis (Thunb.) Klotzsch

6 Garcke, Abh. Konigl. Akad. Wiss. Berlin 1859: 98

(1860). Type: South Africa, Thimberg (UPS-THUNB

11560, holo.).

E. striata var. cuspidata Boiss.: 170 (1862). E. ciis-

pidata Bemh. ex Krauss: 150 (1846), nom. illegit. non.

Bertol. (1843). Type: South Africa, Natal, summit of

Tafelberg, 2 000-3 000’, Sept. 1839, Krauss 441 (MO,

holo.; BM, BOL, M, TCD, iso.). [Krauss (1845) men-

tioned the number "44E, though this did not appear in

Krauss (1846). Boissier (1862) did not cite any speci-

mens and only cited Krauss’ illegitimate name. Conse-

quently the type of Boissier’s name is the same as that

of Krauss’. As for E. kraussiana, the holotype is at MO.]

3. Euphorbia subg. Euphorbia

3a. Sect. Euphorbia

E. aeruginosa Schweickerdt, Bulletin of Miscel-

laneous Information 1935: 205 (1935). Type: South

Africa, Transvaal, Soutpan, Soutpansberg, 1 2 Apr. 1 934,

Schweickerdt & Verdoorn 688 (K, lecto., designated

here; PRE, isolecto.). [Schweickerdt (1935) cited also

‘Soutpan, 23 Nov. 1932, Obermeyer, Schweickerdt &

Verdoorn 15V (PRE) and indicated that both were ‘syn-

types’.]

E. avasmontana Dinter, Sukkulentenforschung in

Siidwestafrika, 11. Teil: 96 (1928). Type: Namibia, near

Windhoek, Auas Mtns, Dinter (PRE, lecto., designated

here). [Although Carter (2002) cited a specimen at B,

this does not exist. A specimen at PRE was annotated by

Dinter himself as "Euph. avasmontana Dtr msc’ . This is

selected as lectotype.]

E. volkmanniae Dinter: 124 (1928). Type: Namibia,

near Otavi, Auros, 1924, Dinter (B, photo).

E. hottentota Marloth: 336 (1930). Type: South

Africa, Cape, Richtersveld, Kubus Kloof, 300 m, 29

Aug. 1925, Marloth 12520 (PRE, lecto., designated

here). [Marloth (1930) also cited Marloth 13357 (mis-

sing).]

E. kalaharica Marloth: 338 (1930). Type: South

Africa, Cape, Neusberg, near Kakamas, 700 m, 15 Aug.

1928, Marloth 14039 (PRE, lecto., designated here).

[Marloth (1930) also cited Marloth 13555 (missing).]

E. sagittaria Marloth: 337 (1930). E. avasmontana

var. sagittaria (Marloth) A.C. White et al.\ 817 (1941).

Type: South Africa, Cape, 12 miles south of Uping-

ton towards Prieska, Aug. 1929, Marloth 14035 (PRE,

lecto., designated here). [Marloth (1930) also cited Mar-

loth 13385 (missing).]

E. venenata Marloth: 337 (1930). Type: Namibia,

Tsarris Mtns, west of Maltahohe, Marloth 4687 (K,

holo.). [Although Carter (2002) cited a specimen at

PRE, this does not exist. Marloth’s description of E.

venenata is vague about such things as the size of the

cyathia and the number of glands in each cyathium.

Nevertheless, the fairly weak spines of the photograph

that he included (figure 7) and the type specimen show

that this is not E. virosa but E. avasmontana.]

The name E. hottentota was maintained as distinct

from E. avasmontana in Bruyns et al. (2006). Marloth

(1930: 335) separated E. avasmontana and E. hotten-

tota by the number of angles on the branches (7-angled

in E. avasmontana', 5-6-angled in E. hottentota) but

White et al. (1941: 824) pointed out that ‘some of Mar-

loth’s herbarium specimens do not agree entirely with

the typical form’ so that the identity of this ‘species’

is less clear than Marloth thought. Over the large area

where it occurs branches are frequently 4-angled and

may have up to eight angles and no clear separation

into 5-6-angled and 7-angled plants is possible. No dif-

ferences in the floral structures have been detected on

which they could be separated.

226

Bothalia 42,2 (2012)

E. barnardii A.C. White et al.. The Succulent Euphor-

bieae 2: 965 (1941). Type: South Africa, Transvaal,

Sekukuniland, farm Driekop, east of Lulu Mountain,

3 000’, 6 Jan. 1937, Barnard 449 (PRE, holo.; MO, iso.).

E. caerulescens Haw., The Philosophical Magazine,

or Annals of Chemistry, Mathematics, Astronomy, Natu-

ral Histoiy and General Science, Ser. 2, 1: 276 (1827).

E. virosa var. caerulescens (Haw.) A. Berger: 81 (1906a).

Type: South Africa, Cape of Good Hope, Bowie, culti-

vated plant at Kew Gardens, pressed Nov. 1876 by N.E.

Brown (K, lecto., designated here).

Possible types for E. caerulescens include (1) a speci-

men ‘Cape of Good Hope, Bowie (K)’, which was made

by N.E. Brown in November 1876 from ‘the type plant

(still in cultivation at Kew) dried by myself’ (Brown

1915: 365) and (2) a drawing by Bond (423/292) of the

apex of a branch and annotated ‘drawn from the plant

from which Haworth described’ and ‘Received in 1823

from the Cape of Good Hope by Mr Bowie’. I propose

that we accept that the plant in cultivation was among

those (if there were more than one) from which Haworth

drew up his description so that I have designated the

specimen made by N.E. Brown as the lectotype.

E. canariensis Thunb.: 86 (1800), nom. illegit., non

L. (1753). Type: South Africa, Thunberg (UPS-THUNB

11416, holo.).

E. /eJ/ert// A. Berger: 80 (1906a). Type: South Africa,

fl. & fr. Aug. 1906, received from collection of F. Ledien

(NY, holo.).

E. ledienii var. dregei N.E.Br.: 366 (1915). Type:

South Africa, near Port Elizabeth, received 9 Sept 1912,

I.L.Drege (K, lecto., designated here). [For E. ledi-

enii var. dregei. Brown (1915) cited two collections:

Humansdorp div., near Zeekoe River, Thunberg-, near

Port Elizabeth, received 9 Sept 1912, I.L.Drege (K). He

annotated both the specimen UPS-THUNB 11416 and

that of Drege as ‘var. dregei' so one is designated as lec-

totype.]

Brown (1915) mentioned that he had not seen any

flowers of E. caerulescens, nor any dried specimens

that he could definitely refer to it, other than the ‘type’.

He distinguished E. caerulescens and E. ledienii by the

glaucous or bluish-green stems, with spines 6-12 mm

long in the former; green, not glaucous stems, with

spines 2-6 mm long in the latter (Brown 1915: 244).

Dyer (1931) and White et al. (1941) found that these

distinctions were not useful and they maintained that the

only difference between E. caerulescens and E. ledienii

was the rhizomatous habit of the former. This charac-

ter was neither mentioned by Haworth nor is it visible

in either the type specimen or the drawing by Bond. It

was also not mentioned by N.E. Brown, who knew the

type specimen in cultivation. Therefore the association

by Dyer (1931) and White et a/.(1941 ) of a rhizomatous

habit with E. caerulescens and a non-rhizomatous habit

with E. ledienii is erroneous and the name E. caerules-

cens must refer to the same non-rhizomatous plants as

E. ledienii. Consequently, E. ledienii is a synonym of

E. caerulescens. This confusion was not recognised in

Bruyns et al. (2006), where E. ledienii was treated as a

separate species from E. caerulescens. The rhizomatous

plants are here treated as a separate species, E. radyeri

Bruyns and the differences between them are discussed

under that species.

E. clavigera N.E.Br, Flora capensis 5(2): 362 (1915).

Type: Swaziland, Bremersdorp (Manzini), 1 800’, 5 Jan.

1905, Burtt-Davy 3010 (K 000253371, holo.; K, PRE,

iso.). [The sheet at K has two specimens of the same

number mounted on it, of which the lower one is anno-

tated as ‘type’. This is therefore designated as holotype.

The specimen at PRE is ‘part of type’.]

E. persistens R. A. Dyer: t. 713 (1938). Type: Moqam-

bique, east of Ressano Garcia, July 1936, F.Z.van der

Merwe E14 sub PRE 23395 (PRE, holo.; K, PRE, iso.).

E. clivicola R. A. Dyer, Bothalia 6: 221 (1951). Type:

South Africa, Transvaal, Lunsklip, 20 miles north of Pot-

gietersrust, 13 Sept. 1946, Plowes sub PRE 28386 (PRE,

holo.; K, iso.).

E. complexa R.A.Dyer, The Flowering Plants of

South Africa 17: t. 643 (1937). Type: South Africa,

Transvaal, road from Louw’s Creek to Kaapmuiden,

June 1936, Van der Merwe 100 sub PRE 21373 (PRE,

holo.; K-2 sheets, W, iso.).

E. confinalis R.A.Dyer, Bothalia 6: 222 (1951). Type:

South Africa, Transvaal, Kruger Nat. Park, 2 miles east

of ‘The Gorge Camp’, 900’, 20 May 1949, Codd & De

Winter 5580 (PRE, holo.; K, NH, iso.).

E. cooperi N.E.Br. ex A. Berger, Sukkulente Euphor-

bien: 83 (1906). Type: South Africa, Natal, Umgeni

Valley, 1862, Cooper, cultivated plant at Kew Gardens,

pressed Sept. 1899 by N.E. Brown (K 00025338, lecto.,

designated here; K, isolecto.). [Brown made two speci-

mens in September 1899 from the plant introduced to

Kew by Cooper in 1862. He labelled both of these ‘Type

specimen’. Leach (1970) selected one of these speci-

mens (though it is not specified which of them) as a neo-

type for E. cooperi. However, although Berger (1906a)

described it from material at La Mortola in Italy, he was

familiar with the plants at Kew and so one of Brown’s

specimens is taken as the lectotype.]

E. eduardoi L.C. Leach, Boletim da sociedade bro-

teriana 42: 161 (1968). Type: Angola, Namibe distr.,

Dois Irmaos, 550 m, 5 May 1960, Mendes 3959 (LISC

011538, holo.; BM, LISC, LUAI, PRE, iso.).

E. enormis N.E.Br., Flora capensis 5(2): 362 (1915).

Type: South Africa, Pietersburg, Sept. 1905, Marloth

5144 (PRE, holo.; K, iso.).

E. excelsa A.C. White et al.. The Succulent Euphor-

bieae 2: 966 (1941). Type: South Africa, Transvaal,

Lydenburg distr., hills near Olifants River, Apr. 1938,

Van der Merwe 1677a (PRE, holo.).

E. grandialata R.A.Dyer, The Flowering Plants of

South Africa 17: t. 641 (1937). Type: South Africa,

Transvaal, Penge mine. Van der Merwe 1002 sub PRE

21372 (PRE, holo.; K, W, iso.).

E. grandicornis A. Blanc, Catalogue and Hints on

Cacti, ed. 2: 68 (1888). Type: Illustration on left hand

Bothalia 42,2 (2012)

227

side of figure on page 68 of A. Blanc, Catalogue & Hints

on Cacti, ed. 2 (1888) (lecto., designated here).

E. grandicornis Goebel: 42, fig. 15 (1889), nom. ille-

git., non A. Blanc ( 1 888).

E. grandicornis J.E. Weiss: 291 (1893), nom. illegit.,

non A. Blanc (1888).

The authorship of this species is usually given as

‘Goebel’ (e.g. Brown (1915); White et al. (1941)) or

‘Goebel ex N.E.Br.’ (e.g. Carter (2002)). However,

while N.E. Brown (1897) published the first detailed

description of E. grandicornis, the name was in use for

a long time before this and there are several earlier brief

descriptions that validated the name. The first known

published appearance of the name E. grandicornis

is Oudemans (1865), but the name was not validly

described there. The earliest validation of the name is

that by A. Blanc ( 1 888), in which it is said that "Euphor-

bia grandicornis is still more remarkable on account

of its tremendous spines and queer, contorted form’.

According to White et al. (1941), a figure of E. gran-

dicornis appeared in an earlier catalogue of A. Blanc of

1 887, but 1 have not been able to trace this. The next one

that has been detected is that of Goebel (1889), in which

the diagnosis is similarly rudimentary but still consti-

tutes valid publication. In J.E. Weiss’ account of 1893

a more detailed diagnosis of E. grandicornis appeared.

Since both Weiss’ and Goebel’s names are illegitimate,

lectotypes are not selected for either of them.

E. grandidens Haw., Philosophical magazine and

journal 66: 33 (1825). Type: Illustration number 807/323

by T. Duncanson at K of specimen received 1822 from

Cape of Good Hope collected by Bowie (lecto., desig-

nated here).

E. evansii Pax: 86 (1909). Type: South Africa, Trans-

vaal, Lowveld, near Barberton, Pole Evans (missing).

[Carter (2002) cited the type specimen at PRE, but this

does not exist, nor is there any material known else-

where that could have been seen by Pax.]

Euphorbia evansii was said to differ (White et al.

1941) from E. grandidens in being shorter (reaching

10 m as opposed to 16 m), with 3- to 4-angled second-

ary branches with gently sinuate margins (as opposed to

3-angled or rarely 2- to 4-angled in E. grandidens with

more prominently toothed margins), spines lacking the

pairs of prickles at their bases, these often present in E.

grandidens. None of these differences are clear-cut and

I have found it impossible to separate the known collec-

tions into two distinct species. Consequently, the name

E. evansii is placed in synonymy, although it was kept

separate in Bruyns et al. (2006).

E. griseola Pax, Botanische Jahrbiicher fur System-

atik 34: 375 (1904). Type: Botswana, Lobatsi, Marloth

3413 (missing). Neotype (Leach 1967): Botswana, 2

miles north of Lobatsi, 16 Jan 1960, Leach & Noel 121

(SRGH, duplicates at BR. G, K, CISC, PRE). [The type

has not been located.]

E. groenewaldii R.A.Dyer, The Flowering Plants

of South Africa 18; t. 714 (1938). Type: South Africa,

Transvaal, 10 miles northeast of Pietersburg towards

Mokeetsi, Nov. 1936, B.H.Groenewald sub Van der

Menve 1186 (sub PRE 23397) (PRE 253379, holo.; K,

PRE, iso.).

E. ingens E.Mey. ex Boiss. in A.P. de Candolle, Pro-

dromus 15(2): 87 (1862). Type: South Africa, Natal,

in woods near Durban, Drege 4614 (S, holo.; K, iso.).

[Boissier (1862) cited a specimen at ‘h. Bunge’ and that

at S was annotated by him, so is taken as the holotype.

That at K is a ‘fragment from type’.]

E. similis A. Berger, Sukk. Euph.: 69 (1906a). Type:

South Africa, Natal ? (missing).

N.E. Brown pressed two specimens from plants in

cultivation at Kew that were reputed to be E. similis

and mentioned that he had sent a branch to Berger who

had confirmed that this was what he named E. simi-

lis. However, many of the pressed branches on the two

specimens at K bear foliage-leaves 15-80 mm long and

consequently they cannot represent either E. ingens

or E. similis in which the leaves were ‘minute’ accord-

ing to Berger and where such foliage-leaves are only

present on the young stem. P.R.O. Bally determined one

of these specimens at K as E. obovalifolia A. Rich. (= E.

ampliphylla Pax) and this is more likely to be the cor-

rect identity of this plant, which Brown (1915) used for

his description of E. similis, but which is not the same as

that which Berger (1906a) described.

E. kaokoensis (A. C. White et al.) L.C. Leach, Dinteria

12: 33 (1976). E. subsalsa var. kaokoensis A. C. White

et al: 965 (1941). Type: Namibia, Kaokoveld, Kauas

Okawe, 28 Nov. 1939, C.J.Hahn sub Otzen 3 (PRE,

holo.).

E. keithii R.A.Dyer, Bothalia 6: 223 (1951). Type:

Swaziland, western edge of Lebombo Mtns, near Stegi,

ft. 1949, Keith sub PRE 28423 (PRE, holo.; GRA, K,

NH, S, SRGH, iso.).

E. knobelii Letty, The Flowering Plants of South

Africa 14; t. 521 (1934). Type: South Africa, Transvaal,

Enselsberg near Zeerust, Sept. 1933, Knobel sub PRE

15854 (K, holo.). [Although Carter (2002) cited the

type from PRE, the specimen is not present there. It is

assumed that this was sent to K on this occasion. This

specimen was collected from the same plant from which

the figure was painted.]

E. knuthii Pax, Botanische Jahrbiicher fur Systema-

tik 34; 83 (1904). Type; Moqambique, Ressano Garcia,

1 000’, 27 Dec. 1897, Schlechter 11949 (K, lecto., des-

ignated here; BM, BOL, BR, G-2 sheets, GRA, HBG,

PRE, WAG, isolecto.). [The sheet at K was annotated

by Pax (‘Knuthii Pax !’) and here he also scratched out

Schlechter’s proposed name for the plant. Nevertheless,

N.E. Brown annotated it as ‘part of type’. This sheet is

then taken as the lectotype. Carter & Leach (2001 ) infor-

mally selected the specimen at K as lectotype, but this is

invalid and so it is formally designated here.]

E. limpopoana L.C. Leach ex S.Carter, Kew Bul-

letin 54: 960 (2000). Type: Zimbabwe, Fulton’s Drift,

25.5 km NNW of Beitbridge, Sept. 1963, Leach 11582a

(SRGH, holo.).

E. malevola subsp. bechuanica L.C. Leach: 6 (1964).

Type: Botswana, halfway between Palapye and Francis-

228

Bothalia 42,2 (2012)

town, Jul. 1937, fl. 1942, Obermeyer (PRE 0645765-0,

holo.; K, PRE, iso.).

E, louwii L.C. Leach, The Journal of South African

Botany 46: 207 (1980). Type: South Africa, Transvaal,

c. 14 km east of Marken, 900 m, 1 Nov. 1975, Leach et

ail 5555 (PRE 0548997-0, holo.; K, PRE, SRGH, iso.)

E. lydenburgensis Schweickerdt & Letty, The Elow-

ering Plants of South Africa 13: t. 486 (1933). Type:

South Africa, Transvaal, Steelpoort Valley, 30 miles

north of Eydenburg, 7 July 1932, Van Balen & De Wyn

sub PRE 14398 (PRE, lecto., designated here; K, iso-

lecto.). [Schweickerdt & Letty (1933) cited two speci-

mens: Van Balen & De Wyn sub PRE 12465 (PRE) and

Van Balen & De Wyn sub PRE 14398 (PRE, K). The lat-

ter is selected as lectotype.]

E. otjingandu Swanepoel, S. African J. Bot. 75: 497

(2009). Type: Namibia, Kunene Region, along Van Zyl’s

Pass 1 km west of Otjihende, 1 305 m, 1 May 2007,

Swanepoel 268 (WIND, holo.; PRU, iso.).

E. otjipembana L.C. Leach, Dinteria 12: 29 (1976).

Type: Namibia, north of Otjipemba, Leach & Can-

ned 15044 (PRE, holo.; BM, K, LISC, M, MO, SRGH,

WIND, iso.).

E. perangusta R. A. Dyer, The Flowering Plants of

South Africa 18: t. 716 (1938). Type: South Africa,

Transvaal, Koedoesrant, north of Zeerust, Jan. 1936,

Louw 99 {sub PRE 23399){VWE, holo.; BOL, GRA, K-2

sheets, MO, P, SRGH, iso.).

E. pseudocactus A. Berger, Sukkulente Euphorbien:

78 (1906). Type: Country unknown, but probably India,

branch from the type plant, received from A. Berger Oct.

1910 (K, lecto., designated here).

Euphorbia radyeri Bruyns, sp. nov., a E. caerules-

cente caulibus crassioribus, plus profunde articulatis,

exterioribus rhizomatosis differt. Type: South Africa,

Cape, 20 miles from Kendrew towards Jansenville, Jan.

1930, Dyer 2357 (GRA, holo.; PRE, iso.).

Bisexual spiny glabrous succulent shrub 1-2 m tall,

1-3 m broad, branching extensively mainly from base

of similar main stem with woody and fibrous roots, with

many peripheral branches spreading underground from

plant for up to 0.5 m by rhizomes and then rising erect

from soil. Branches 30-70 mm thick, strongly con-

stricted into many ± spherical segments, smooth, grey-

green; tubercles fused into 3-7 wing-like often sinuate

angles, laterally flattened and rounded and projecting

3-10 mm from angles, spine-shields around apex and

united into continuous homy and later somewhat corky

brown to grey or black margin, 4-6 mm broad in upper

part tapering to 2-3 mm below, bearing 2 spreading and

widely diverging brown to grey spines (2-)6-15 mm

long; leaf-rudiments on tips of new tubercles towards

apex of branches and main stem, 1-4 mm long, 2-4 mm

broad, spreading, fleeting, broadly ovate, obtuse, sessile,

with green-brown obtuse ± pyramidal stipule on either

side at base. Inflorescences in large numbers per branch

towards apex, each a group of 1-3 cymes in axil of

tubercle, on peduncle 2-4(6) mm long, 2—3 mm thick,

each cyme with 3 vertically disposed cyathia, central

male, outer 2 female only (or bisexual) and developing

later, with 2 ovate bracts 1.0-1. 5 mm long and 1. 5-2.0

mm broad subtending cyathia; cyathia cupular-conical,

glabrous, 3. 5-6.0 mm broad (2-3 mm long below inser-

tion of glands), with 5 lobes with deeply incised mar-

gins, bright yellow; glands (3-)5, transversely oblong

to kidney-shaped or rectangular, 2-3 mm broad, bright

yellow, ascending-spreading, slightly convex to concave

above, outer margins entire and slightly raised; stamens

entirely glabrous, bracteoles palmate and enveloping

groups of stamens, deeply and finely divided, glabrous;

ovary globose, glabrous, included to slightly exserted

on erect pedicel 1. 5-2.0 mm long and soon becoming

slightly exserted, calyx slightly extended around base;

styles 2-4 mm long, branched in upper third. Capsule

6-7 mm diam., obtusely 3-angled, glabrous, erect and

exserted on short pedicel 2-4 mm long.

Although E. caerulescens and E. radyeri are similar,

they are easily separated. Branches around the perimeter

of most plants of E. radyeri are usually rhizomatous and

this phenomenon is unknown in E. caerulescens. The

branches tend to have a more bluish green colour in E.

radyeri than in E. caerulescens, though the colour var-

ies greatly in the latter, with greener branches on plants

from more sheltered habitats. The branches of E. rady-

eri are thicker, deeply articulated into almost spherical

segments, while those of E. caerulescens are generally

more slender and only indistinctly articulated into con-

siderably longer, cylindrical segments. In E. radyeri the

tubercles are often much longer and broader and the

leaf-rudiments are somewhat larger than in E. caerules-

cens. Florally E. caerulescens and E. radyeri are very

similar. In E. caerulescens the cyathia are often slightly

narrower, beeoming more abruptly narrow beneath the

glands, while the female florets are borne on a slightly

longer pedicel and are without the elongated calyx of E.

radyeri.

E. restricta R. A. Dyer, Bothalia 6: 224 (1951). Type:

South Africa, Transvaal, The Downs, 4 500’, 14 Oct.

1947, Codd & De Winter 3092 (PRE 0248764-0, holo.;

GRA, K-2 sheets, NH, PRE-2 sheets, SRGH, iso.).

E. rowlandii R. A. Dyer, Bothalia 7: 28 (1958). Type:

South Africa, Transvaal, Kruger Nat. Park, 8 miles north

of Punda Maria, 1 600’, 25 July 1951, Rowland Jones

48 (PRE 0248767-0, holo.; K-2 sheets, PRE, SRGH-2

sheets, iso.).

E. schinzii Pax, Bulletin Herbier Boissier 6: 739

(1898). Type: South Africa, Transvaal, Berea Ridge,

Barberton, 3 100’, 13 Feb. 1891, Galpin 1297 (BOL,

lecto., designated here; K, isolecto.). Pax (1898) also

cited ‘South Africa, Transvaal, Pretoria, Rehmann 4347’

(missing).

E. sekukuniensis R. A. Dyer, The Flowering Plants

of South Africa 20: t. 775 (1940). Type: South Africa,

Transvaal, Steelpoort River, north of Roossenekal, Aug.

1938, Van der Merw’e 1765 (sub PRE 25475) (PRE

0248772-1, holo.; GRA, PRE, SRGH, iso.).

E. stellata Willd., Species Plantarum 2: 886 (1799).

Type: Illustration in F. le Vaillant, Reise Itin. Ed. Germ.

Francof 4: 245, t. 1 1 (1797) (lecto., designated here).

Bothalia42,2 (2012)

229

E. procumbens Meerburgh: t. 55 (1789), nom. illegit.,

non Mill. (1786).

E. radiata Thunb.: 86 (1800). Type; South Africa,

Cape, (UPS-THUNB 11547, holo.).

E. uncinata DC.: 151 (1805). Type: Illustration in DC

(1805) by Redoute opposite p. 151 (lecto., designated

here). [De Candolle (1805) did not cite any specimens

and none annotated as E. uncinata by him have been

found.]

E. squarrosa Haw.: 276 (1827). Type: Illustration

number 295/423 by G. Bond at K of specimen from

Cape of Good Hope (lecto., designated here). [No type

was designated by Haworth (1827) nor, in this case,

did he refer to a collection of Bowie. There is a speci-

men at Kew made by N.E. Brown soon after he arrived

at Kew in 1873. This was from a very old plant which

was ‘believed to have been introduced by Bowie and

so may have been one of the original plants from which

Haworth described the species’. Since there is some

uncertainty surrounding whether Haworth saw this

specimen, the drawing number 295/423 by G. Bond is

selected as lectotype.]

E. micracantha Boiss.: 25 (1860). Type: South Africa,

Cape, between Zuurberg and Klein Bruintjieshoogte,

2 ()00-2 500’, Oct. Drege 8206a (K, lecto., designated

here; MO, S, isolecto.). [Boissier (1860) cited ‘inter

Zuurebergen et Klein Bruintjeshoogte et inter Vis-

chrivier et Fort Beaufort {Drege n° 8206)\ The col-

lection from ‘between Fish R. & Fort Beaufort’ is now

labelled Drege 8206c (K) and the other as Drege 8206a

(K, MO, S).]

E. gilbertii A.Berger; 39 (1906a). Type: South Africa,

Cape, Cooper (missing).

E. lombardensis Nel: 194 (1933b). Type: South

Africa, Cape, Mortimer, 1 200-1 300 m, Dec. 1933,

M.Lombardsub SUG 1564 (NBG).

White et al. (1941) recognised three species: E. mic-

racantha (plants with mainly 4-angled, erect branches,

low tubercles less than 4 mm long and relatively long

spines), E. squarrosa (plants with mainly 3 -angled, often

spreading branches, particularly prominent tubercles

4-8 mm long and relatively short spines) and E. stel-

lata (plants with mainly 2-angled, spreading branches

usually pressed to the ground, relatively low tubercles

less than 4 mm long and relatively short spines). How-

ever, they illustrated many plants which were intermedi-

ate between these three and expressed doubt that three

species could be distinguished: ‘And in the event that

distinct species are involved, their limits can hardly be

defined accurately’ (p. 730). This arrangement of three

species was followed in Bruyns et al. (2006). However,

it is quite often impossible to place a plant with certainty

under one of these three names and so a broader view is

taken here and a single species is recognised.

E. subsalsa subsp. fluvialis L.C. Leach, Dinteria 12:

29 (1976). Type: Angola, Ruacana Falls, Leach & Can-

ned 14509 (FISC, holo.; BM, K, LUAI, M, MO, PRF,

SRGH, iso.).

E. tetragona Haw., The Philosophical Magazine, or

Annals of Chemistry, Mathematics, Astronomy, Natural

History and General Science Ser. 2,1: 276 (1827). Type:

Illustration number 291/1060 by G. Bond at K of speci-

men received in 1823 from Cape of Good Hope col-

lected by Bowie (lecto., designated here). [There are two

paintings of E. tetragona by Bond and this one, where

details of the cyathia are shown, is selected as the lecto-

type.]

E. tortirama R. A. Dyer, The Flowering Plants of

South Africa 17: t. 644 (1937). Type: South Africa,

Transvaal, Bandolierskop, Sod & S. W.Smith sub PRE

21371 (PRE 0258980-1, holo.; K, PRE, W, iso.).

E. triangularis Desf. ex A.Berger, Sukkulente

Euphorbien: 57 (1906). Type: South Africa, Cape, cul-

tivated plant at Kew Gardens, pressed 30 Oct. 1913 by

N.E. Brown (K, lecto., designated by Dyer 1974b).

E. umfoloziensis Peckover, Aloe 28: 37 (1991).

Type: South Africa, Natal, near Dingaanstat, 10 Apr.

1981, Peckover (PRE, holo.).

E. vandermerwei R.A.Dyer, The Flowering Plants

of South Africa 17: t. 660 (1937). Type: South Africa,

Transvaal, White River, Sept. 1936, Van der Mem’e sub

PRE 22436 (PRE, holo.; K-2 sheets, P, SRGH, iso.).

[The specimens at P, SRGH and one at K lack the PRE

number but are ‘from Type Specimen’ so are taken as

isotypes as well.]

E. venteri L.C. Leach ex R. Archer & S. Carter, The

Flowering Plants of Africa 57: 86 (2001). Type: Bot-

swana, near Tsessebe, c. 45 km north of Francistown,

12 Dec. 1991, Venter et al. 174 (PRE, holo.; K, UNIN,

iso.).

E. virosa Widd., Species Plantarum 2: 882 (1799).

Type: Illustration in Paterson, Reisen: 60, t. 9, 10 (1790)

(lecto., designated here). [These two figures were cited

by Willdenow (1799) and are considered here to con-

stitute a single plate, suitable as a lectotype. This figure

was cited by Leach (1971) and Carter (2002), but in nei-

ther case was it formally designated as lectotype.]

E. bedica Hiem: 945 (1900). Type: Angola,

Moqamedes distr., frequent in sandy coastal hills from

Giraul up to Cape Negro, Jul. 1859, Welwitsch 643 (BM,

holo.).

E. dinteri A.Berger: 109 (1906b). Type: Namibia,

Khan River, received 1904, Dinter (NY, holo.). [The

specimen in the Alwyn Berger Herbarium consists of

seeds only. These are annotated by Berger as follows:

‘11069, von C. Dinter als E. virosa eingefiihrt. 1904’.

They are therefore the seeds which Berger (1906b) men-

tioned, that had been sent to him by Dinter. Their large

size makes it clear that they came from plants of E.

virosa.}

E. virosa f. caespitosa H. Jacobsen: 81 (1955). Type:

none cited.

E. virosa f striata H. Jacobsen: 81 (1955). Type: none

cited.

230

Bothalia 42,2 (2012)

E. waterbergensis R.A.Dyer, The Flowering Plants

of Africa 28: t. 1095 (1951). Type: South Africa, Trans-

vaal, 2.5 miles north of Elmerston P.O. towards Ellisras,

3 300’, Apr. 1948, Codd & Erem 4018 (PRE 0248809-0,

lecto., designated here; BOL, K, PRE, SRGH, isolecto.).

[There are two sheets of this at PRE, neither annotated

as ‘Type’ and so the present one is selected as lectotype.]

E. zoutpansbergensis R.A.Dyer, The Flowering

Plants of South Africa 18: t. 715 (1938). Type: South

Africa, Transvaal, Wylliespoort, Sept. 1937, Dyer 3873

sub PRE 23393 (PRE 0248810-0, holo.; E, K, MO,

PRE-2 sheets, US, iso.).

3b. Sect. Monadenium (Pax) Bruyns

E. lugardiae (N.E.Br.) Bruyns, Taxon 55: 413 (2006).

Monadenium lugardiae N.E.Br.: 138 (1909). Type:

Botswana, foot of Kwebe Peak, Kwebe Hills, 3 500’,

fl. Aug. 1897 & leaves Feb. 1898, Mrs Lugard 22 (K,

holo.).

3c. Sect. Tirucalli Boiss.

E. gummifera Boiss., Centuria Euphorbiarum: 26

(1860). Type: South Africa, Cape, low-lying areas

between Verleptpram and the mouth of the Orange

River, Sept. 1830, Drege 2944 (P, holo.; S, iso.).

E. gregaria Marloth, Transactions of the Royal Soci-

ety of South Africa 2: 36 (1910). Type: Namibia, Kuibis,

Marloth 4683 (PRE, holo.; K, iso.).

E. congestiflora L.C. Leach, Boletim da sociedade

broteriana, ser. 2, 44: 197 (1970). Type: Angola, Namibe

distr., between Cumilunga & Curoca Rivers, 11 Jan.

1956, Mendes 1265 (LISC, holo.; BM, LUA, M, SRGH,

iso.).

E. damarana L.C. Leach, Bothalia 11: 500 (1975).

Type: Namibia, Damaraland, c. 64 km west of Khorixas,

27 July 1973, Leach & Canned 15064a (LISC, holo.; K,

M, PRE, SRGH, WIND, iso.). [Although Leach (1975b)

stated that the holotype is at PRE, it is at LISC.]

4. Euphorbia subg. Rhizanthium (Boiss.) Wheeler

E. albipollinifera L.C. Leach, South African Journal

of Botany 51: 281 (1985). Type: South Africa, Cape,

Springbokvlakte, Dec. 1978, Bruyns 1826 (NBG, holo.;

K, PRE, iso.).

E. arida N.E.Br., Flora capensis 5(2): 319 (1915).

Type: South Africa, Cape, Britstown div., near De Aar,

Schonland (K, holo.).

E. benthamii Lliern, Catalogue of the African plants

collected by Dr. Friedrich Welwitsch in 1853-61, 1: 943

(1900). Type: Angola, between Lopollo and Ivantala,

Feb. 1860, Welwitsch 283 (BM, holo.; K, LlSU, iso.).

E. brakdamensis N.E.Br., Flora capensis 5(2): 324

(1915). Type: South Africa, Cape, Brakdam, 1 600’, 7

Sept. 1897, Schlechter 11123 (K, holo.; BOL, BR, GRA,

IIBG, L-2 sheets, PRE, S, WAG, iso.).

In Bruyns et al. (2006) E. brakdamensis was included

under E. filiflora. Careful examination of Schlechter’s

many pressings of the type collection of E. brakda-

mensis, shows, however, that this is not correct. In E.

filiflora the stem and branches are very similar in shape

and thickness, with the stem usually slightly longer than

the branches, if it can be detected at all. In E. brakda-

mensis, on the other hand, the branches are very much

more slender than the stem, which is largely buried

in the ground and is greatly exceeded in height by the

branches. E. flliflora has unusually long cyathia (often

around 8 mm long), with especially long styles (7-9 mm

long) and long male pedicels. The cyathia in E. brakda-

mensis do not exceed 5 mm long and the styles are not

longer than 6 mm. The marginal processes on the glands

in E. brakdamensis are much more brightly coloured

than those of E. flliflora where, however, they are longer,

more slender and considerably more numerous.

E. braunsii N.E.Br., Flora capensis 5(2): 326 (1915).

Type: South Africa, Cape, Aberdeen distr., without pre-

cise locality, Brauns (K, holo.). [Although Brown (1915)

cited two specimens, he mentioned, in addition, that the

species was described from the collection of Brauns and

so the Brauns collection at K is taken as the holotype.]

E. rudis N.E.Br.: 322 (1915). Type: Namibia, sandy

plains northeast of Narudas Slid, 28 Dec. 1912, Pearson

8141 (BOL, lecto., designated here; SAM, isolecto.).

[Of the collections cited by Brown (1915) only Pearson

4310 (BOL, K) and Pearson 8141 (BOL, SAM) have

duplicates and so Pearson 8141 is selected as the lecto-

type.]

E. marientalii Dinter: 31 (1914). Type: Namibia,

Mariental, Dinter 3164 (SAM, holo.).

E. rangeana Dinter: 31 (1914). Type: none cited.

[Euphorbia rangeana was very similar to E. marientalii

and was distinguished by 'E. rangeana ist griinbraun

und graubraun’ (Dinter 1914: 31), which does make it

validly published. However, no specimens were cited

here.]

Euphorbia rudis was maintained as a distinct ‘spe-

cies’ in Bruyns et al. (2006). However, for E. rudis and

E. braunsii White et al. (1941: 474) mentioned that

‘there is really no sharp line of distinction between the

two plants, but rather a gradation. The typical forms of

the two are fairly clearly distinguishable, while many

of the intermediate forms are very confusing indeed

and difficult to classify satisfactorily.’ The distinctions

between the two included: the smaller ‘average size of

the main stem’, the ‘more slender’ branches with the

tubercles ‘somewhat more recurved at the apex’ and

‘rather smaller’ cyathia and ‘more completely united

styles’ in E. rudis. These are all subject to considerable

variation so that the name E. rudis has been abandoned

here.

E. brevirama N.E.Br., Flora capensis 5(2): 317

(1915). Type: South Africa, Cape, Jansenville div., near

Klipplaat, Schonland 1716 (K, holo.). [Carter (2002)

listed a specimen at GRA but this does not exist.]

E. bruynsii L.C. Leach, The Journal of South Afri-

can Botany 47: 103 (1981). Type: South Africa, Cape,

Steytlerville, Bruyns 1814 (PRE, holo.; SRGH, iso.).

Bothalia 42,2 (2012)

231

E. bubalina Boiss., Centuria Euphorbiarum: 26

(1860). Type: South Africa, Cape, among thorn-bushes

near Buffelsrivier, Drege 4615 (P, holo.). [Boissier

(1860) cited a specimen in ‘h. Bunge’, so this sheet is

taken as the holotype.]

E. laxiflora Kuntze: 286 (1898). Type: South Africa,

East London, 5 Mar. 1894, Kuntze (NY, holo.; K, iso.).

E. bupleurifolia Jacq., Plantarum rariorum horti cae-

sari schoenbrunnensis descriptiones et icones 1: 55, t.

106 (1797). Tithymalus bupleunfolius (Jacq.) Haw.: 138

(1812). Type: Illustration in Jacq., PI. Hort. Schdnbr. 1:

t. 106 (1797) (lecto., designated here).

E. proteifolia Boiss.: 92 (1862). Type: South Africa,

near Umtata. Drege 8196 (missing). [Boissier (1862)

cited a specimen in ‘h. Bunge’, but this has not been

located.]

E. caperonioides R. A. Dyer & P.G.Mey., Mitteilun-

gen aus der Botanischen Staatssammlung Miinchen 6:

245 (1966). Type: Namibia, Kaokoland, 3 miles west of

Etanga, 7 Apr. 1957, De Winter & Leistner 5420 (PRE,

holo.).

E. caput-medusae L., Species Plantarum 1: 452

(1753). Type: J. Burm., Rar. Afric. PL: t. 8 (1738)

(lecto., designated by Wijnands 1983).

E. fructus-pini Mill.: Euphorbia no. 10 (1768). Medu-

sea fructus-pini (Mill.) Haw.: 134 (1812). Neotype (des-

ignated here): J. Burm., Rar. Afric. PL: t. 8 (1738).

When Miller (1768) ‘described’ E. fructus-pini, he

referred to Linnaeus (1737) and Boerhaave (1720: 258).

He also referred to it as ‘Euphorbium Afrum facie fruc-

tus pini’ and then added ‘African Euphorbium with the

appearance of Pine fruit, commonly called Little Medu-

sa’s Head’. In the longer discussion after the literature

citations, he added ‘The tenth sort hath a thick short

stalk, which seldom rises more than eight or ten inches

high, from which come out a great number of trail-

ing branches which are slender, and grow about a foot

in length; these intermix with each other like those of

the seventh sort, but they are much smaller, and do not

grow near so long, but have the same appearance, from

whence it is called Little Medusa’s Head: the ends of

these branches are beset with narrow leaves, between

which the flowers come out, which are white, and

shaped like those of the other species.’

Linnaeus (1737) referred to Boerhaave (1720: 258)

and ‘Breyne, Prodr. 2: 100’. In Boerhaave (1720: 258)

one finds ‘8. ..in capitis Medusae’ and ‘9. Euphorbium;

Afrum; facie fructus pini ’....Tithymalus, Africanus,

arborescens, squamato caule, spinosis MH 3:344’. ‘MH

3’ refers to the third volume of ‘Planta Historia universa-

lis’ (Morison 1699). On page 344 of this work, Morison

referred to ‘Pluk. Phyt. t. 230’. In Plukenet (1692), the

phrase ‘Tithymalus, Africanus, arborescens, squamato

caule, spinosis’ appears under t. 230, fig. 5 as well as

‘pini fructu facie’. This figure is of E. loricata.

Euphorbia loricata does not produce trailing

branches from a ‘thick short stalk’, nor is it ‘without

spines, having tubercles furnished with very narrow

leaves’. Therefore Miller’s information makes it clear

that his name cannot be applied to E. loricata, even

though some of the references he gave refer to that spe-

cies. The reference to ‘very narrow leaves’ makes it

more likely that this name refers to E. caput-medusae

than E. inermis, among the species with a ‘thick short

stalk’ and ‘trailing branches’. At present no preserved

material of Miller’s Euphorbia no 10 is known and so a

neotype is selected.

E. caput-medusae var. geminata Alton: 136 (1789).

Type: Illustration in J. Burm., Rar. Afric. PL: t. 9, fig. 1

(1738) (lecto., designated here).

E. caput-medusae var. major Alton: 135 (1789).

Type: Illustration in Commelijn, Praeludia Bot.: t. 7

(1703) (lecto., designated here).

E. caput-medusae var. minor Alton: 135 (1789).

Type: Illustration in Breyne, Prodr. rar. pi. sec.: t. 19

( 1 739) (lecto., designated here).

E. tuberculata Jacq.: 43, t. 208 (1797). Dacty’lanthes

tuberculata (Jacq.) Haw.: 133 (1812). Medusea tuber-

culata (Jacq.) Klotzsch & Garcke: 61 (1860). Type:

Illustration in Jacq., PL Hort. Schonbr. 2: t. 208 (1797)

(lecto., designated here).

E. medusae Thunb.: 86 (1800). Type: South Africa,

Cape, Thunberg (UPS-THUNB 11494, lecto., designated

here). [Thunberg (1800) placed two of his collections

under E. medusae, namely UPS-THUNB 11494 and

1 1495. The latter is a piece of E. hamata.]

Medusea major Haw.: 134 (1812). Type: Illustration

in Commelijn, Praeludia Bot.: t. 7 (1703) (lecto., desig-

nated here).

Medusea tessellata Haw.: 135 (1812). E. tessellata

(Haw.) Sweet: 107 (1818). Type: none cited.

E. commelinii DC.: 110 (1813). Type: Illustration in

Commelijn, Praeludia Bot.: t. 7 (1703) (lecto., desig-

nated by Wijnands 1983).

E. fructus-pini var. geminata Sweet: 356 (1826).

Type: Illustration in J.Burm., Rar. Afric. PL: t. 9, fig. 1

( 1 738) (lecto., designated here).

E. bolusii N.E.Br.: 333 (1915). Type: South Africa,

Transvaal, near Middelburg?, Sept. 1886, H. Bolus 9767

(BOL, holo.; K, iso.). [The locality given is considered

to be an error (White et al. 1941 : 372).]

E. ramiglans N.E.Br.: 306 (1915). Type: South

Africa, Namaqualand, 1883, H. Bolus sub BOL 9448

(BOL, holo.; K, iso.).

E. marlothiana N.E.Br.: 331 (1915). Type: South

Africa, Cape, near Neu Eisleben, fl. Oct. -Nov. 1914,

Marloth 5733 (PRE, holo.; BOL, NBG, K, iso.).

E. muirii N.E.Br.: 331 (1915). Type: South Africa,

Cape, Platbos, Still Bay, Muir 174 (BOL, lecto., desig-

nated here; PRE, SAM, isolecto.). [Brown (1915) also

cited the following: Albertinia, Muir (K), Pearson sub

SAM 2261 (K, SAM).]

232

Bothalia 42,2 (2012)

E. tiiberculatoides N.E.Br.: 332 (1915). Type: South

Africa, Cape, Theefontein, Malmesbury div., Bachmann

1042 (K, lecto., designated here). [Brown (1915) also

cited the following: Grey (K), Bolus 4359 (BOL).]

E. ?uacowanii N.E.Br.: 334 (1915). E. tuberculata var.

macowanii (N.E.Br.) A.C. White et al.\ 372 ( 1941 ). Type:

South Africa, Clanwilliam [wrongly labelled as Cannon

Hill, Uitenhage], MacOwan 3286 (K, lecto., designated

here; SAM, WU, isolecto.). [Brown (1915) also cited the

following: Schlechter 8419 (GRA, K, PRE).]

E. confhiem Nel: 193 (1933b). Type: South Africa,

Cape, open flats, Kliphoogte, Sept. 1929, Herre

sub SUG 5549 (missing). Type: Illustration in Kak-

teenkunde: 194 (1933) (lecto., designated here).

[Although Carter (2002) cited a specimen at STE (now

incorportated into NBG), this does not exist.]

E. celata R.A.Dyer, Bothalia 11: 278 (1974). Type:

South Africa, Vanrhynsdorp distr., Moedverloor, 100 m,

12 May 1973, Hall 4272 (PRE, holo.).

E. miscella L.C. Leach: 341 (1984a). Type: South

Africa, Cape, near Lekkersing, Leach et ah 16545 (NBG,

holo.; PRE, iso.).

E. clandestina Jacq., Plantarum rariorum horti cae-

sari schoenbrunnensis descriptiones et icones 4: 43, t.

484 (1804). Type: Illustration in Jacq., PI. Hort. Schonbr.

4: t. 484 (1804) (lecto., designated here).

E. clava Jacq., Icones plantarum rariorum 1 (4): 9,

t. 85 (1784). Treisia clava (Jacq.) Haw.: 131 (1812).

Type: Jacq., Icon. 1: t. 85 (1781) (lecto., designated by

Wijnands 1983).

E. canaliculata Lam.: 417 (1788). Type: South

Africa, collector unknown (P-LAM P00381883, holo.).

[A specimen of this ‘species’ is present in the Lamarck

herbarium at P and is taken as the holotype and

Wijnands’ lectotype (Wijnands 1983: 99) is set aside.]

E. coronata Thunb.: 86 (1800). Type: South Africa,

Cape, Thunberg (UPS-THUNB 11434, holo.). Treisia

tuberculata Haw.: 65 (1819). Type: Introduced by D.

Young to Epsom, 1815, fl. Chelsea 1818 (missing).

E. pubiglans N.E.Br.: 338 (1915). Type: South Africa,

Cape, near Port Elizabeth, Sept. 1912, l.L.Drege (K,

holo.).

E. clavarioides Boiss., Centuria Euphorbiarum:

25 (1860). Type: South Africa, Cape, Sneeuberge at

Poortjie, Drege 8200 (P; duplicates at K, S, W) (lecto.,

designated here). [Boissier (1860) did not state which

herbarium he saw this collection in. This suggests that

there was a specimen at G, but this has not been located.

A leetotype is selected.]

E. clavarioides var. truncata (N.E.Br.) A.C. White et

ai: 309 (1941). £. truncata N.E.Br.: 309 (1915). Type:

South Africa, Standerton, Burtt-Davy 1953 (K, lecto.,

designated here). [Brown (1915) also cited the fol-

lowing: Transvaal, 23 Nov. 1905, Leendertz 670 (K);

Leendertz 1873 (K) and WHms 1339 (missing); Kolbe

(BOL).]

E. basutica Marloth: 408 (1910a). Type: Lesotho,

Leribe, Dieterlin (eult. Phillips, fl. Cape Town in Mar.

1909) sub Marloth 4671 (K, holo.; NH, PRE, SAM,

iso.). [Though Marloth (1910a) cited no number, it is

assumed that this is the same specimen as his type.]

E. colliculina A.C. White et al., The Suceulent

Euphorbieae 2: 962 (1941). Type: South Africa, Cape,

2.5 miles north of Oudtshoom, Aug. 1939, Dyer 4053

(PRE 0247438-2, leeto., designated here; BOL, K,

PRE, isolecto.). [According to Dyer’s collecting book

the number should be 4053 not 4052, as given in White

et al (1941); the latter has no entry next to it while the

former is 'E. colliculina WDS sp. nov. type’. White et

al (1941) designated Dyer 4052 the ‘type’ and Marloth

10577 (K, PRE) the ‘type of capsule’ so a lectotype is

designated here.]

In Bruyns et al (2006), E. colliculina was included

under E. esculenta. However, while they bear a close

resemblance to one another, there are many differences

and two distinct species are involved. Mature speci-

mens of E. colliculina are altogether more delicate than

those of E. esculenta and neither the main stem nor the

branches reach the thickness that are normal for E. escu-

lenta. E. esculenta also produees several swollen roots

from the base of the tap-root and this phenomenon is

unknown in E. colliculina, where the thick taproot tapers

off quite abruptly into slender, fibrous roots. Florally E.

colliculina is also easily separated from E. esculenta in

that the cyathial lobes and the bracteoles within the cyat-

hium laek the densely bushy hairiness at their apices that

make the cyathium of E. esculenta distinctively furry or

woolly. The cyathial glands are also much larger than

those of E. esculenta.

E. crassipes Marloth, Transactions of the Royal Soci-

ety of South Africa 1:318 (1909). Type: South Africa,

Cape, Biesiespoort, Marloth (4399)4397 (PRE, holo.; K,

iso.).

E. fusca Marloth: 38 (1910b). Type: South Africa,

Cape, Britstown, Sept. 1909, Marloth 4682 (PRE, holo.;

K, iso.). [In the cases of both E. crassipes and E. fusca.

Brown annotated the specimens at K as parts from Mar-

loth’s type specimens and so the holotype is the speci-

men at PRE in each case, with isotypes at K.]

E. baliola N.E.Br.: 327 (1915). Type: Namibia, Great

Karas Mountains, between 1st & 2nd outspan between

Kraikluft and Narudas Slid, 5400’, 26 Dec. 1912, Pear-

son 8095 (K, holo.; BOL-2 sheets, GRA, SAM, iso.).

[Brown annotated the specimen at K himself as ‘Type’

but those at BOL and SAM were not annotated by him.

Therefore that at K is taken as the holotype.]

E. inornata N.E.Br.: 586 (1925). E. inelegans

N.E.Br.: 322 (1915), nom. illegit., non N.E.Br. (1911).

Type: South Africa, Cape, near Kimberley, Sept. 1912,

Moran {sub Schonland 1718) (K 000253322, holo.;

GRA, K, iso.). [Brown (1915) mentioned that E. inor-

nata was described from a living plant sent by Schon-

land in 1912, grown at Kew and pressed by Brown

himself in June 1913. This is the specimen ‘near Kim-

berley, Moran, living plant sent to Kew by Schon-

land’ (K000253322). Mounted on the same sheet is

Bothalia 42,2 (2012)

233

another specimen, namely ^ Moran sub Schonland 1718'

(K000253323) and both had ‘Type’ written on them by

Brown. The former is taken as the holotype.]

E. eendoornensis Dinter: 196 (1932). Type: Namibia,

between Wittsand and Eendom, 26 Mar. 1924, Dinter

(missing). Neotype (designated here): Namibia, Vrede,

Briiyns 11362 (NBG).

E. hopetownensis Nel: 192 (1933b). Type: South

Africa, Cape, Hopetown, 1930, E.Markoetter sub SUG

5529 (missing). Type: Illustration in Kakteenkunde: 192

(1933) (lecto., designated here). [Although Carter (2002)

cited a specimen at STE (now incorportated into NBG),

this does not exist.]

Marloth (1910b) said that Euphorbia fusca differed

from E. crassipes by the non-persistent peduncles (some

peduncles being persistent in E. crassipes). In most pop-

ulations of E. crassipes one finds plants with persistent

peduneles and others without them so this character can-

not be used to distinguish between them and the type of

E. crassipes at PR£ is a typical specimen of what is usu-

ally referred to as "E. fusca’. White et al. (1941) main-

tained that the main differences between E. crassipes

and E. fusca were the slightly more cylindrical stem,

thicker branches, the deeper involucres and the green

glands. However, in the description Marloth did not

mention the glands at all and they were only represented

in a small black and white drawing so that their colour

was unknown. None of these other differences are sig-

nificant in this widely distributed and quite variable spe-

cies.

Although the glands of E. inornata were given as

olive-green on their upper surface, which is unusually

pale for E. crassipes, the shape of the plant, the relative

thickness of the branches and the shape of the cyathia

and glands all fit E. crassipes, under which it is included

here.

Euphorbia hopetownensis was described from a small

plant (only 5 cm broad) with ascending, relatively stout

branches which bore unusually short peduncles at 5-7

mm long and ‘pink-purple’ glands with five teeth. The

small figure in the text and these few details are strongly

suggestive of E. crassipes, under which this name is

subsumed here.

Euphorbia baliola was not listed in Bruyns et al.

(2006) but is included here under E. crassipes. Brown

(1915) believed it to differ from E. crassipes by the dif-

ferent manner in which the tubercles on the stem are

formed (from the persistent bases of the branches), the

presence of branches right to the centre of the stem and

the longer pedicels of the male florets with longer hairs.

However, collections made near where the type was col-

lected are typical of E. crassipes except for somewhat

more slender branches and it seems improbable that two

distinct species are involved here.

E. crotonoides Boiss. in A.P. de Candolle, Prodromus

15(2): 98 (1862). Type: Sudan, Kordofan, near El Obeid

in shade of Adansonia, Kotsehy 419 (S, holo., K, iso.).

[Boissier (1862) cited a specimen in ‘h. Vindob.’ and

since the sheet at S is from ‘Herb. Musei Palat. Vindob.’,

this is taken as the holotype.]

E. cumulata R. A. Dyer, Records of the Albany

Museum 4: 92 (1931). Type: South Africa, Cape, Botha

Ridge, 10 miles from Grahamstown on Queen’s Road,

Dyer 669 (GRA, holo.; K, iso.).

E. cylindrica Marloth ex A. C. White et al.. The Suc-

culent Euphorbieae 2: 962 (1941). Type: South Africa,

Cape, Kubiskow Mtn, 7 Sept. 1926, Marloth 12860

(PRE, holo.).

E. davyi Pax ex N.E.Br., Flora capensis 5(2): 305

(1915). Type: South Africa, Transvaal, near Pretoria, 19

Nov. 1901, J.W.C. Kirk 48 (K, lecto., designated here;

PRE, isolecto.). [Brown (1915) listed three specimens:

Kirk 48 (K, PRE), Burtt-Davy 2196 (K) and Burtt-Davy’

5562 (K) and annotated the first and last as ‘Type’, so a

lectotype is selected. At K there is a letter written by Pax

from Breslau in Feb. 1906 to Burtt-Davy that requested

his permission to name this species after him.]

E. pseudohypogaea Dinter: 265 (1921a). Type:

Namibia, am Wege von Oas nach Gobabis, Dinter 3144

(missing).

E. bergii A.C. White et al.: 963 (1941). Type: South

Africa, Orange Free State, Koffiefontein, Scholtz (miss-

ing).

E. pseudoduseimata A.C. White et al.: 963 (1941).

Type: Namibia, Hohenhorst, 45 miles SW of Wind-

hoek. Nov. 1940, Otzen (PRE, holo.; K, iso.). [White

et al. (1941) cited a specimen from ’45 miles SW of

Windhoek, Otzen 37’ as the type, and mentioned the

farm-name ‘Hohenhorst’ (p. 414) as well. At PRE there

is a specimen ‘Hohenhorst, SW of Windhoek, Nov.

1940, Otzen PRE 4 5881'. The material at Kew lacks

the number ‘37’, but is from the same locality and was

annotated by Dyer as ‘Part of type specimen’. Therefore

the specimen at PRE is taken as the holotype.]

E. decepta N.E.Br., Flora capensis 5(2): 320 (1915).

Type: South Africa, Cape, near Willowmore, Brauns

1712 {K, holo.).

E. albertensis N.E.Br.: 323 (1915). Type: South

Africa, Cape, near Prince Albert, between railway and

village near Prince Albert, May 1907, Marloth 4397 (K,

holo.; PRE, iso.).

E. astrophora Marx: 311 (1996). Type: South Africa,

Cape, north of Klipplaat, Marx 204 (GRA, holo.).

E. gamkensis Marx: 38 (1999a). Type: South Africa,

Cape, south of Calitzdorp, Marx 225 (GRA, holo.).

E. suppressa Marx: 33 (1999a). Type: South Africa,

Cape, near Seekoeigat, Marx 227 (GRA, holo.).

According to N.E. Brown (1915), Marloth considered

his number 4397 from between Prince Albert and Prince

Albert Road (‘the railway’) to belong to E. crassipes.

However, Brown believed that the absence of a ‘flat

top to the stem’ was significant and that it represented

a distinct species, which he named E. albertensis. The

plants pressed (K, PRE) have a relatively slender stem

(far too slender to belong to E. crassipes) with numerous

short branches towards their apex (which are also much

234

Bothalia 42,2 (2012)

more slender than in E. crassipes) with many long,

slender, spine-like persistent, sterile peduncles. Vegeta-

tively these plants are extremely similar to E. decepta

and, although Brown (1915) was unable to supply much

detail about the floral parts of E. albertensis, this name

is included here under E. decepta.

Both E. gamkemis and E. suppressa were treated as

distinct species in Bruyns et al. (2006). but are here rel-

egated to synonymy.

Euphorbia suppressa was compared extensively with

E. albertensis and E. arida (Marx 1999a). The basis for

comparison with E. albertensis was mainly Figure 445

of White et al. (1941). However, it is uncertain whether

this figure is of the ‘species’ described by N.E. Brown

as E. albertensis. Apart from the fact that Dyer (GRA

records) had tentatively attributed two specimens from

the area between Prince Albert and Klaarstroom to E.

arida, it remains unclear what this new species has to

do with E. arida (a species of the north-eastern Great

Karoo and southern Free State) and why it was not com-

pared with E. decepta, which is fairly well-known on the

southern portion of the Great Karoo between Beaufort

West and Willowmore. Florally E. arida and E. decepta

are not easily separated except by the somewhat shal-

lower cyathium (and slightly shorter styles) with fewer,

often obsolete teeth on the outer margins of the cyat-

hial glands in E. decepta (deeper cyathium, longer style

and more prominent and more numerous marginal teeth

in E. arida). However, although the plant appears to be

very similar in both species, beneath the soil plants of

E. arida have a system of swollen tuberous roots which

develop from and extend the tap-root. These structures

are entirely absent in E. decepta. In all these respects E.

suppressa is identical to E. decepta and so this name is

included here under E. decepta.

Euphorbia gamkensis was compared extensively

with E. crassipes (and its synonym E. fusca). However,

it differs from E. crassipes by its much smaller stature

(main stem at most 90 mm thick) by the considerably

deeper cyathium whose glands are more-or-less without

marginal processes (these are particularly prominent in

E. crassipes and are usually strongly deflexed). Again,

it ought to have been considered how it differs from E.

decepta. Vegetatively the two are difficult to separate

and I have been unable to find any reliable differences.

The cyathia differ in that the styles are shorter and more

deeply divided in E. gamkensis, but no other significant

differences have been detected. As I consider this to be

insufficient on which to base a separate and otherwise so

similar species, I have included E. gamkensis under E.

decepta.

While E. astrophora was compared with many spe-

cies, including E. decepta (Marx 1996), it was said that

it ‘very closely resembles’ E. decepta, differing by the

slightly shorter branches and the convex glands. The

glands may be concave in E. decepta as well, and plants

of E. decepta are very variable in size so that there are

no substantial differences between them. There are

therefore no grounds for separating E. astrophora from

E. decepta.

E. dregeana E.Mey. ex Boiss. in A.P. de Candolle,

Prodromus 15(2): 95 (1862). Type: South Africa, Cape,

between Koussie and Silverfontein in Kaus Mtn, 2 000’,

29 Aug. 1830, Drege 2942 (P, holo.; G, K-2 sheets, S,

iso.). [Boissier (1862) cited a specimen in ‘h. Bunge’, so

the specimen in P is taken as the holotype. A specimen at

MO is excluded as it is unnumbered and has been anno-

tated ‘must be 2942’, for which no grounds are known.]

E. duseimata R. A. Dyer. t. 530 (1934). Type: Bot-

swana, ± 100 miles northwest of Molepolole, flowered

in cultivation in Pretoria in Nov. 1931, G.J.de Wyn sub

72426 (PRE, holo.).

E. esculenta Marloth, Transactions of the Royal

Society of South Africa 1: 319 (1909). Type: South

Africa, Cape, Klipplaat (Graaff-Reinet), received living

Sept. 1907, Marloth 4162 (PRE, holo.; BOL, K, SAM,

iso.).

E. fasciculata Thunb., Prodromus plantarum capen-

sium 2: 86 (1800). Type: South Africa, Cape, Thunberg

(UPS-THUNB 11456, holo.).

E. filiflora Marloth, Transactions of the Royal Soci-

ety of South Africa 3: 123 (1913). Type: South Africa,

Cape, near Concordia, Apr. 1912, Krapohl sub Marloth

5119 (NBG, lecto., designated here; K, PRE, isolecto.).

[Marloth (1913) mentioned two collections, one made

by himself at Chamis in Great Namaqualand (i.e. south-

ern Namibia) in October 1910 and another sent to him

from Concordia in Namaqualand (i.e. in north-western

South Africa) by Krapohl in March 1912. He appears to

have recorded both of these under his number 5119 but

E. ferox Marloth, Transactions of the Royal Society

of South Africa 3: 122 (1913). Type: South Africa, Cape,

Klipplaat, 1905, Marloth 5147 (PRE, holo.; BOL, iso.).

E. alternicolor N.E.Br.: 344 (1915). E. aggregata

var. alternicolor (N.E.Br.) A. C. White et al.\ 616 (1941).

Type: South Africa, N.S.Pillans (K, holo.).

E. captiosa N.E.Br.: 345 (1915). Type: South Africa,

Cape, near Aberdeen, flow. Sept. 1904, Schonland 1661

(GRA, holo.).

E. elastica Marloth: 37 (1910b), nom. illegit., non

Poisson & Pax (1902). Type: South Africa, Cape, near

Anenous, Nov. 1908, Carstens sub Marloth 4684 (PRE,

holo.).

E. ecklonii (Klotzsch & Garcke) BailL, Adanso-

nia 3: 144 (1863). Tithymalus ecklonii Klotzsch &

Garcke: 68 (1860). Type: South Africa, Cape of Good

Hope, Swellendam district, Breede River at Swellen-

dam (70.10), hills under 1000’, Aug., Ecklon & Zeyher,

Euphorb. 16 (W, holo.; P, S, iso.).

E. pistiifolia Boiss.: 93 (1862). Type: South Africa,

Cape of Good Hope, Swellendam district, Breede River

at Swellendam (70.10), hills under 1 000’, Aug., Ecklon

& Zeyher, Euphorb. 76 (S, lecto., designated here; P, W,

isolecto.). [Boissier (1862) also cited Drege 8195 (S,

W), which is from the same locality.]

E. inermis var. laniglans N.E.Br.: 328 (1915). Type:

South Africa, Cape, near Klipplaat, received Oct. 1912,

Marloth 5270 (K, holo.; PRE, iso.).

Bothalia42,2 (2012)

235

I have not been able to locate any material of the col-

lection from Chamis. He wrote ‘Type’ on the specimen

at NBG and not on any of the other pieces distributed

under this number.]

E. fUiflora var. nana G.Will.: 49 (2003). Type: South

Africa, Cape, T’Gabies Plateau, northwest of Kosies,

Oct. 1999, Williamson 5933 (BOL, holo.).

E. nelii A. C. White et al.: 484 (1941). E. meyeri Nel:

134 (1933a), nom. il legit, non Steud. (1840) nec Boiss.

(1860). Type: South Africa, Cape, Klipfontein, c. 1 000

m. Sept. 1929, Herre sub SUG 5545 (missing). Type:

Illustration in Kakteenkunde: 134 (1933) (lecto., desig-

nated here). [Since the type of E. meyeri Nel and conse-

quently of E. nelii White et al. is missing, a lectotype is

selected.]

E. versicolores G.Will.: 284 (1995). Type: South

Africa, Cape, near Eksteenfontein, Williamson 4453

(NBG, holo.).

E. flanaganii N.E.Br., Flora capensis 5(2): 314

(1915). Type: South Africa, Cape, grassy slopes near

Kei Mouth, 100’, June 1893, Flanagan 1800 (PRE

0254449-0, holo.; K, PRE, iso.). [The specimen at K is

annotated ‘branches from the Type Specimen (in Cape

Town Herb.)’. There is no type specimen of this species

in any Herbarium in Cape Town but there are two sheets

at PRE, one of which is annotated by Brown as ‘Type’.

This is taken as the holotype.]

E. ernestii N.E.Br.: 307 (1915). Type: South Africa,

Cape, Hospital Hill, near Queenstown, 3 600’, 17 Sept.

1911, Galpin 8066 (K 000253285, holo.; K, PRE, iso.).

E. gatbergensis N.E.Br.: 310 (1915). Type:

South Africa, Cape, near Gatberg (south of Elliott),

3 000-3 500’, Baur 251 (K, holo.).

E. franksiae N.E.Br.: 315 (1915). Type: South Africa,

Natal, Camperdown, 2 000’, 19 Oct. 1910, Franks sub

Medley-Wood 11727 (K, holo.; NH, PRE, iso.). [The

specimens at NH and PRE were not seen by Brown,

though that at NH is annotated as ‘part of Type Spec.’.

The sheet at K contains two specimens, one collected

by Franks on 19 October 1910, pressed by Wood and

sent to K (this being the other ‘part of Type Spec.’) and

another made from two plants sent in Apr. 1913 to, and

cultivated at, Kew. Only the former specimen is anno-

tated by Brown as ‘type’ and is taken as the holotype.]

E. woodii N.E.Br.: 315 (1915). Type: South Africa,

Natal, Clairmont Flats, Wood 4090 (K, lecto., designated

here; NH, isolecto). [Brown (1915) cited also: Clairmont

Flats, Wood 11803 (K) and Wood 12612 (K).]

E. passa N.E.Br.: 313 (1915). Type: South Africa,

Natal, Cooper, cult. J. Corduroy, 6 July 1905 (K

000253311, lecto., designated here, K, isolecto.).

[Brown (1915) cited also: Scottsburg, Pole Evans (miss-

ing); Umzumbi, Wood(K).]

E. discreta N.E.Br.: 316 (1915). Type: South Africa,

Natal, banks of Umzimkulu River near shore, 25 Feb.

1837, Bachmann 757 (K, holo.).

Brown (1915) recognised a host of ‘species’ here,

including E. discreta, E. ernestii, E. flanaganii, E. frank-

siae, E. gatbergensis, E. passa, and E. woodii. White

et al. (1941) reduced the number slightly by placing E.

discreta and E. passa in synonymy under E. woodii and

recognising E. ernestii, E. flanaganii, E. franksiae, E.

gatbergensis, and E. woodii as distinct species. Brown

(1915: 314) commented on the remarkable extent to

which these plants can vary in size; in particular, how

one of them increased in size in cultivation from 30^0

branches at 3-8 inches long to 140 branches that were

9-14.5 inches long and this underlines the vegetative

variability that one may observe here. Nevertheless, he

distinguished E. flanaganii from E. woodii by the ‘much

shorter branches’ (Brown (1915): 314) and E. discreta

from E. woodii by the fact that the ‘body of the plant is

much smaller’ (Brown (1915): 316). As commented on

extensively by White et al. (1941), this makes no sense

in view of such strong variation in the size of individu-

als. Plants producing more than one rosette of branches

are not unusual and are found in many populations.

Although this feature was not mentioned by Brown

(1915) in his descriptions, this was supposed to separate

E. gatbergensis from E. ernestii (White et al, 1941: 75),

but they recognised that plants of both ‘species’ could

produce several rosettes. E. flanaganii and E. woodii

were separated by ‘Ovary puberulous = E. fanaganiP;

Ovary glabrous to thinly pubescent with long hairs = E.

woodii' (White et al. (1941): 75, adapted from Brown

(1915): 239). In practise some populations have plants

with pubescent ovaries and others with glabrous ovaries

and to distinguish two species on the basis of the length

and density of this pubescence is untenable. Conse-

quently all these names are reduced here to synonymy

under a single species.

E. fortuita A. C. White et al.. The Succulent Euphor-

bieae 2: 962 (1941). Type: South Africa, Cape, 27 miles

from Ladismith towards Barrydale, Aug. 1939, Dyer

4074 (PRE, Sheet I, holo.; K, PRE-2 sheets, iso.).

Euphorbia fortuita was included under E. esculenta

in Bruyns et al. (2006). However, although in both spe-

cies the cyathial glands are mostly dark and the centre

of the cyathium is densely filled with white hairs, there

are significant differences between them that warrant

their recognition as distinct species. In E. fortuita the

glands are much broader and the cyathium is more coni-

cal, having a rather rounded, almost spherical shape in

E. esculenta. Furthermore, the pedicels of the male flo-

rets in E. esculenta are glabrous (densely pubescent in E.

fortuita) but in E. fortuita the bracteoles are uniformly

pubescent in their upper half, while in E. esculenta they

are densely pubescent only at their apices. The ovary is

entirely glabrous in E. esculenta and densely pubescent

above in E. fortuita.

E. friedrichiae Dinter, Neue und wenig bekannte

Pflanzen Deutsch-SWA’s: 29 (1914). Type: Namibia,

Warmbad, comm. Sept. 1913, M. Friedrich sub Dinter

3253 (SAM, holo.).

E. gariepina Boiss., Centuria Euphorbiarum: 28

(1860). Type: South Africa, Cape, Verleptpram, interior

at Orange River, Drege 8214 (G, holo.; K, S, W, iso.).

236

Bothalia 42,2 (2012)

E. gariepina subsp. balsamea (We/w. ex Hiern)

L.C. Leach, Excelsa Taxonomic Series 2: 78 (1980). E.

balsamea Welw. ex Hiem: 951 (1900). Type: Angola,

Welwitsch 634 (K, holo.; G, P, iso.).

E. bergeriana Dinter: 28 (1914). Type: Namibia,

Okawayo near Karibib, Dinter 1385 (SAM, holo.).

E. schaeferi Dinter: 304 (1921b). Type: Namibia,

Klein Karas, Schafer sub Dinter 1233 (SAM, lecto.,

designated here). [Dinter (1921b) eited: Holoog, Dinter

1233; Klein Karas, Schafer. Since Holoog is close to

Klein Karas, it is assumed that the specimen cited here

is one of these, although the details do not quite corre-

spond.]

E. gerstneriana Bruyns, nom. nov.

E. franksiae var. ziiluensis A. C. White et ah: 962

(1941). Type: South Afriea, Natal, near Mahlabatini, 18

Oct. 1935, Gerstner 687 (PRE, holo.).

E. gerstneriana is closely allied to E.flanaganii. In E.

flanaganii, the branches form a dense, usually strongly

spreading erown around the apex, which is itself devoid

of branehes. This bare apex of the stem is green with

prominent tubercles and is somewhat depressed towards

the eentre. In E. gerstneriana the branches are produeed

right to the apex of the stem so that the apex of the

stem is not visible at all. The branehes in E. flanaganii

are usually distinctly swollen towards their bases while

in E. gerstneriana the branehes are uniformly thick to

their bases. They are also much less densely clustered

around the apex of the stem and form an aseending, usu-

ally lax rosette. The cyathia differ in that they are pale

green and distinctly red-veined on the lobes and in the

subtending braets in E. gerstneriana, with deep brown-

ish purple, comparatively small glands that are widely

spaced around the eyathium. In E. flanaganii the cyathia

and their subtending bracts are yellow-green, the glands

are usually bright yellow and are far broader, usually

almost eontiguous around the eyathium. The styles of

E. gerstneriana are particularly broad (more than twice

the breadth of those in E. flanaganii) and form an almost

mushroom-like top to the female floret.

E. globosa (Haw.) Sims, Curtis’ Botanical Magazine

53: t. 2624 (1826). Dactylanthes globosa Haw.: 382

( 1 823). Medusea globosa (Haw.) Klotzsch & Garcke: 61

(1860). Type: Illustration number 808/15 by T. Duncan-

son at K of specimen received 1821 from Cape of Good

Hope collected by Bowie (lecto., designated here).

E. glomerata A. Berger: 104 (1906a). Type: South

Africa, Cape (missing).

E. hallii R.A.Dyer, The Journal of South Afriean Bot-

any 19: 135 (1953). Type: South Africa, Cape, Botter-

kloof. May 1953, Hall sub PRE 28532 (PRE, holo.;

GRA, K, iso.).

E. hamata (Haw.) Sweet, Hortus suburbanus Lond-

inensis: 107 (1818). Medusea hamata (Haw.) Klotzsch

& Garcke: 251 (1859). Dactylanthes hamata Haw.: 133

(1812). Type: Illustration in J.Burm., Ran Afric. PI.: t.

6, figure 3 (1738) (lecto., designated here). [This fig-

ure was cited by Haworth (1812). It was cited by Carter

(2002) as ‘T: ieono’ but this does not constitute valid

leetotypification.]

E. cervicornis Boiss.: 27 (1860). Type: South Africa,

Cape, Heerenlogenment, Zeyher 1530 (G, leeto., desig-

nated here; BOE, SAM, isolecto.). [Boissier (1860) also

cited Drege 2950 (missing).]

E. peltigera E.Mey. ex Boiss.: 91 (1862). Type: South

Africa, Cape, on roeks at Orange River near Verlept-

pram, 19 Sept. 1830, Drege 2951 (S, lecto., designated

here; K, isolecto.). [Boissier (1862) cited a specimen ‘in

h. Bunge’. The specimen in S may be that formerly in

Bunge’s herbarium and could be the holotype but this is

not certain and so it is chosen as leetotype. A sheet at

MO, ‘assumed to be 295 1 ’ is excluded.]

E. heptagona L., Species Plantarum 1: 450 (1753).

Type: Illustration in Boerh., Ind. Alter. Hort. Eugd.-Bat.

1: figure opposite p. 258 (1720) (leeto., designated here).

[This figure was eited by Linnaeus (1753). It was also

cited by Carter (2002) as ‘T: icono’ but this does not

constitute valid leetotypification. Jarvis (2007) stated

that it remained untypified.]

Anthacantha desmetiana Lem.: 64 (1858). Type:

South Africa, Cape, cult. L. Desmet (missing).

E. enopla Boiss.: 27 (1860). Type: South Africa,

Cape, Witpoortsberg, 2 000-3 000’, Aug., Drege 8207

(S, holo.; BM-2 sheets, K, MO, P, W-2 sheets, iso.).

[Boissier (1860) did not cite a herbarium here and so

a leetotype is ehosen The speeimen at MO is a mixed

sheet of which only the left hand and middle pieees are

this species.]

E. heptagona var. /w/vApmo A.Berger: 109 (1902b).

Type: none cited.

E. morinii A.Berger: 98 (1906a). Type: South Africa,

Cape, cultivated material sold by Co. Haage & Schmidt-

Erfurt (missing).

E. atrispina N.E.Br.: 342 (1915). Type: South Afriea,

Cape, near Prinee Albert, received 1912, Pearson (K,

holo.).

E. heptagona var. dentata (A.Berger) N.E.Br.: 351

(1915). E. enopla var. dentata A.Berger: 95 (1906a).

Type: South Africa, Cape, Witpoortsberge, Drege (P,

leeto., designated here). [Berger (1906a) did not state

where the specimen was that he saw, so a leetotype is

designated.]

E. heptagona var. ramosa A. C. White et ai: 964

(1941). Type: South Afriea, Cape, 17 miles north of

Oudtshoorn, Aug. 1939, Dyer 4049 (PRE, holo.; GRA,

iso.).

E. heptagona var. subsessilis A. C. White et al.: 964

(1941). Type: South Africa, Cape, 17 miles east of Ladi-

smith (15 miles west of Calitzdorp), Aug. 1939, Dver

4067 (PRE, holo.).

E. heptagona var. viridis A. C. White et al.: 964

(1941). Type: South Africa, Cape, 1 1 miles west of Cal-

itzdorp in Huis River Pass, Aug. 1939, Dyer 4065 (PRE,

holo.).

Bothalia 42,2 (2012)

237

E. enopla var. viridis A. C. White et al.: 964 (1941).

Type: South Africa, Cape, 17 miles north of Jansenville

towards Graaff-Reinet. Aug. 1939, Dyer 4008 (PRE,

holo.).

E. atrispina var. viridis A. C. White et al.: 964 (1941).

Type: South Africa, Cape, 12-15 miles from Montagu

near Ouberg Pass, Aug. 1939, Dyer 4094 (PRE, holo.).

E. huttonae A.£'.5r., Flora capensis 5(2): 316 (1915).

E. inermis var. huttonae (N.E.Br.) A. C. White et al.: 395

(1941). Type: South Africa, Cape, Carlisle Bridge, on

the Fish River, fl. Nov. 1903, H. Hutton (K, holo.; GRA,

iso.). [N.E. Brown based his description on a small

dried specimen sent to Kew by Schonland in June 1913.

Brown kept two branches and one ‘flower’ at Kew and

sent one branch back to GRA. Thus, although he anno-

tated each as ‘Half of the type specimen’, that at Kew is

actually two thirds of the specimen and is taken as the

holotype.]

E. superans Nel ex Herre: 15 (1950). Type: South

Africa, Eastern Cape, July 1948, Rosenbrock sub SUG

7215 (missing). Neotype (designated here): South

Africa, Carlisle Bridge, Nov. 1903, H. Hutton (GRA,

holo.; duplicate at K). [The specimen cited here by

Herre is missing (though Carter (2002) cited it as being

at STE, now incorporated into NBG). The name is usu-

ally cited as ^E. superans Nel’ but the article in which it

was published was written by H. Herre. No photograph

was included with the protologue and so a neotype is

selected.]

Euphorbia huttonae is re-instated at the level of spe-

cies for various reasons. Vegetatively it differs from E.

inermis in that the rootstock does not develop a series

of swollen, fusiform roots below the stem, but tapers

rapidly off into fine roots. There are several clear dif-

ferences in the cyathia. In E. huttonae the whole of the

upper surface of the gland is bright yellow. Each gland

may be divided deeply down the middle into two broad,

convex, yellow structures which remain pressed together

towards their bases or it may be an entire, solid wedge-

shaped structure that is convex above. The outer edges

of the glands are irregularly toothed and notched and

may be slightly paler in some populations. In E. inermis

each gland possesses a dark green part towards the base

above which it is divided deeply and finely into antler-

like, white processes. Other floral differences are the

spreading, white cyathial lobes in E. inermis (rather than

the pale yellowish green inwardly pressed lobes of E.

huttonae) and the longer styles in E. inermis which are

only divided near their apex (divided much more deeply

to near their middle in E. huttonae).

Some confusion exists over the identity of Euphorbia

superans, which was maintained as a distinct species in

Bruyns et al. (2006). A figure appeared in the Euphor-

bia Journal (Vol. 2: 138, as ‘supemans’) which was cited

by Carter (2002) as E. superans, but the slender, bright

green branches and finely toothed, broad cyathial glands

make it clear that this figure is of E. fianaganii. Herre

(1950) compared E. superans with E. inermis and men-

tioned that the glands were ‘yellow... shortly bifid with

two processes denticulate at the apex, divided [to] about

a third with two diverging processes. ..slightly revolute’.

This is very similar to the structure of the glands in E.

huttonae but is not similar at all to that in E. fianaga-

nii. The length of the styles and the length to which they

are divided also correspond closely to E. huttonae under

which E. superans is now included.

E. hypogaea Marloth, Transactions of the Royal

Society of South Africa 2: 37 (1910). Type: South

Africa, Cape, on the Nieuweveld near Beaufort West,

1 300 m, Nov. 1908, Marloth 4692 (PRE, holo.; K, iso.).

E. inermis Mill., The Gardener’s Dictionary, ed. 8:

Euphorbia no. 13 (1768). Neoype (designated here):

South Africa, Cape, near Swartkops R. and on hills near

Addo, Zeyher 1098 (K; duplicate at SAM,). [Miller

(1768) cited no material and none is known to exist from

this date. Therefore a neotype has been selected.]

E. insarmentosa P.G.Mey., Mitteilungen aus der Bot-

anischen Staatssammlung Miinchen 6: 246 (1966). Type:

Namibia, Outjo distr., Welwitschia, 19 Mar. 1967, Giess,

Volk & Bleissner 6128 (M, holo.).

E. jansenvillensis Nel, Jahrbuch der Deutschen

Kakteen-Gesellschaft 1: 32 (1935). Type: South Africa,

Cape, near Jansenville, Apr. 1932, Le Roux sub SUG

6550 (missing). Neoype (designated here): South Africa,

Cape, 1.5 miles east of Jansenville, Dyer 4012 (PRE).

[Although Carter (2002) cited a specimen at STE (now

incorportated into NBG), this does not exist.]

E. tubiglans Marloth ex R. A. Dyer: 268 (1935). Type:

South Africa, Cape, near Steytlerville, Aug. 1929, Herre

1596 (K, holo.; PRE, iso.).

E. lignosa Marloth, Transactions of the Royal Soci-

ety of South Africa 1: 316 (1909). Type: Namibia, near

Tschaukaib, 400 m, Nov. 1908, Marloth 4637 (PRE,

holo.; BOL, K, iso.). [Brown wrote ‘Part of Type’ on a

specimen of Marloth 5070 at K, but this is incorrect.]

E. engleriana Dinter: 263 (1921a). Type: Namibia,

zwischen Ababis und Habis, Apr. 1913, Dinter 2815

(SAM, holo.).

E. curocana L.C. Leach: 111 (1975a). Type: Angola,

± 18 km southeast of Cumilunga, Mendes 1260 (CISC,

holo.; BM, COI, LUAI, iso.).

E. loricata Lam., Encyclopedie methodique 2(2):

416 (1788). Type: Illustration in Pluk., Phytographia

3: t. 230, figure 5 (1692) (lecto., designated here).

[Lamarck (1788) also cited ‘Petiver Gaz., t. 86, fig. 519’

and ‘Buc’hoz, Dec. 9, t. 3’, but of these three, Pluke-

net’s figure appeared first. These figures all appear to be

copies (sometimes modified by the author) of the fig-

ure, assumed to be by Heinrich Claudius, that is among

the collection of paintings made during the expedi-

tion of Simon van der Stel to the Copper Mountains of

Namaqualand in 1685-6 and known as the Codex Witse-

nii (Wilson et al. 2002). The original figure of Claudius

appears to have been unknown to Lamarck. There is no

specimen of this species in Lamarck’s herbarium at R]

E. hystrix Jacq.: 43, t. 207 (1797). Treisia hystrix

(Jacq.) Haw.: 131 (1812). Type: Illustration in Jacq., PI.

Hort. Schonbr 2: t. 207 (1797) (lecto., designated here).

238

Bothalia 42,2 (2012)

E. armata Thunb.: 86 (1800). Type: South Africa,

Cape, Thiwberg (UPS-TUUNB 11412, holo.).

E. eustacei N.E.Br.: 122 (1913). Type: South Africa,

Cape, near Matjiesfontein, Oct. 1912, C.E. Pillans (K

000253356, holo.; K, PRE, iso.). [From the material sent

by Pillans and cultivated at Kew, N.E. Brown made and

annotated three specimens on two sheets at K and also

sent ‘part of the type’ to PRE. Brown annotated only one

of them (K 000253356) as ‘Type Specimen’ (others as

‘Type, branches from type plant’ and ‘Type Plant’) and

so this is taken as the holotype and the others as iso-

types.]

In Bruyns et al. (2006) E. eustacei was maintained as

distinct from E. loricata. This does not reflect the posi-

tion correctly. Dense, low-growing and mound-forming

plants with slightly broader, more obovate leaves and

spines drying out white have always been taken as typi-

cal of E. eustacei and were assumed to be restricted to

the Matjiesfontein area (White et al 1941), while the

more diffuse, taller plants with narrower leaves and

spines drying out brown that are characteristic of the

valley of the Olifants River between Citrusdal and

Clanwiliam are typical of E. loricata. Nevertheless,

White et al. (1941) hinted at a wider distribution for

E. loricata and included some more densely branched

plants (e.g. figure 264) in their concept of this species.

Now that the respective distributions have become better

known it has been found that there is a gradation from

the one into the other as one progresses eastwards from

the valley of the Olifants River to the Great Escarpment

(rather than two disjunct and distinct species each con-

fined to particular areas) so that E. eustacei and E. lori-

cata are ecotypes of one considerably more widespread

species.

E. maleolens E. Phillips, The Flowering Plants of

South Africa 12: t. 459 (1932). Type: South Africa, near

Bandolierskop, Dec. 1925, C.A. Smith sub PRE 8465

(PRE, holo.).

E. mammillaris L., Species Plantarum 1: 451 (1753).

Type: Commelijn, Praeludia Bot.: t. 9 (1703) (lecto.,

designated by Wijnands 1983).

E. fimbri at a Scopoli: 8 (1788). Type: Illustration in

Delic. FI. Faun. Insubr. 3: 8, t. 4 (1788) (lecto., desig-

nated here).

E. emieagona Haw.: 184 (1803). Type: none located.

E. erosa Willd.: 27 (1814). [Willdenow (1814) gave a

description but cited no specimens.]

E. scopoliana Steud.: 615 (1840), uom. superfl. [Steu-

del (1840) believed that E. fimbriata Scopoli was illegit-

imate, with the name used earlier by Raeuschel, but no

such name has been traced.]

E. mammillaris var. spinosior h.BQXgQx: 109 (1902b).

Type: South Africa, Cape, probably ex hort. F. Ledien

(missing).

E. mammillaris var. submammillaris A. Berger: 1 25

(1902c). Type: South Africa, Cape, cultivated plant from

Berlin Botanic Garden (missing).

E. latimammillaris Croizat: 331 (1933). Type: none

cited.

E. platymammillaris Croizat: 333 (1933). Type: none

cited.

E, matabelensis Pax, Annalen des K. Naturhistor-

ischen Hofmuseums 15: 51 (1900). Type: Zimbabwe

(Matabeleland), Peuther 944 (W, holo.; BM, iso.).

E. currorii N.E.Br.: 545 (1911). Type: Angola, Ele-

phant’s Bay, Curror 29 (K, holo.).

E. ohiva Swanepoel: 249 (2009). Type: Namibia,

Kaokoveld, Hartmann Valley above Cunene River, 470

m, 12 Jan. 2006, Swanepoel 250 (WIND, holo.; PRE,

iso.).

E. melanohydrata Nel, Jahrbuch der Deutschen

Kakteen-Gesellschaft 1: 31 (1935). Type: South Africa,

Cape, flats at Swartwater, Oct. 1930, Pierre sub SUG

6533 (missing). Neotype (designated here): South

Africa, 4 km east of Beesbank, March 1985, Williamson

3401 (BOL). [Although Carter (2002) cited the type at

STE, now incorportated into NBG, no material of the

type has been located and a neotype is selected of mate-

rial from the same area where the type originated.]

E. meloformis Aiton, Hortus Kewensis, ed. 1,2: 135

(1789). Type: Illustration by F. Masson at BM of speci-

men introduced 1774 from Cape of Good Hope col-

lected by Masson (lecto., designated here).

E. pomiformis Thunb.: 86 (1800). E. meloformis var.

pomiformis (Thunb.) Marloth: 45 (1928). Type: South

Africa, Zwartkops, Thunberg (missing).

E. falsa N.E.Br.: 586 (1925). E. meloformis subsp.

meloformis f. falsa (N.E.Br.) Marx: 32 (1999b). E.

infausta N.E.Br.: 358 (1915), nom. illegit., non N.E.Br.

(1912). Type: South Africa, Cape, sheet 332, specimen

annotated ‘dead plant-split-1810’ by Haworth (OXF)

(lecto., designated here). [Brown (1915) cited two speci-

mens: South Africa, without locality, N.S. Pillans sub

BOL 10684 (BOL) and ‘Herb. Haworth’. The latter is

designated as lectotype.]

E. pyriformis N.E.Br.: 359 (1915). Type: cultivated

plant at Kew of unknown origin, pressed by N.E. Brown

14 Jan. 1913 (K, holo.).

E. valida N.E.Br.: 356 (1915). E. meloformis subsp.

valida (N.E.Br.) G.D.Rowley: 97 (1998).' Type: South

Africa, Cape, Jansenville div., near Waterford, received

26 Aug. 1912, l.L. Drege (K, holo.).

E. meloformis var. prolifera Frick: 74 (1934). Type:

Cultivated material from seed imported from South

Africa, A.C.S. 5- 1 1 2-006 (missing).

E. meloformis subsp. meloformis f. magna R. A. Dyer

ex Marx: 13 (1999b). Type: South Africa, Cape, Kwa

Ncwane, Peddie (3327AA), 18 Mar. 1999, Marx 550

(GRA, holo.).

E. monteiroi Hook.f, Bot. Mag. 91: t. 5534 (1865).

Type: Angola, Monteiro (K, holo.).

Bothalia 42,2 (2012)

239

E. marlothii Pax: 36 (1889). Type; Namibia, Karibib,

1 000 m, May 1886, Marloth 1425 (PRE, lecto., des-

ignated here). [The number of the type collection was

given as 4425 by Pax (1889), but this is assumed to be

an error. There is no evidence that Pax saw this speci-

men so it is designated as a lectotype.]

E. longibracteata Pax: 742 (1898). Type: Namibia,

Rehoboth, 1892, Fleck 447a (Z, holo.).

E. baiimii Pax: 636 (1908). Type: Angola, left bank

of Cubango River above Kui marva, 1 100 m, 23 Nov.

1899, Baum 458 (Z, holo.).

E. monteiroi subsp. ramosa L.C. Leach, Kirkia 6:

138 (1968a). Type: South Africa, Transvaal, 10 miles

south of Mica, Leach 11999 (PRE, holo.; BM, BOL,

COI, G, K, LISC, M, SRGH, WIND, Z, ZSS, iso.).

E. monteiroi subsp. brandbergensis Nordenstam,

Dinteria 11; 23 (1974). Type: Namibia, Brandberg,

between Tsisab and Konigstein, c. 1 750 m, 29 May

1963, Nordenstam 2786 (S, holo.).

E. multiceps A. Berger, Monatsschrift flir Kak-

teenkunde 15: 182 (1905). Type: South Africa, Cape,

Karoo near Matjiesfontein, 950 m, Marloth 3450 (miss-

ing). Type: South Africa (NY, lecto., designated here).

[The specimen in the Herbarium of Alwyn Berger at

NY consists of several small stems but has no informa-

tion apart from the name on it. It was undoubtedly seen

by Berger and may well be the Marloth specimen, but is

designated here as lectotype.]

E. multifolia A.C. White et al.. The Succulent Euphor-

bieae 2; 962 (1941). Type: South Africa, Cape, 30 miles

from Laingsburg towards Ladismith, Aug. 1939, Herre

(PRE, lecto., designated here). [White et al. (1941)

listed two specimens from the same locality, collected

by Smith and Herre respectively and designated that

by Smith the type. This being missing, the specimen of

Herre is designated as lectotype.]

E. namaquensis N.E.Br., Flora capensis 5(2): 325

(1915). Type: South Africa, between Aggeneys and

Pella, Pearson 2992 (BOL, lecto., designated by Wil-

liamson 2007; K, SAM, isolecto.).

E. multiramosa Nel: 29 (1935). Type: South Africa,

Cape, Little Bushmanland, flats between Jakkalswater

and Vioolsdrift, Oct. 1930, Herre sub SUG 5890 (miss-

ing). Neotype (designated here): South Africa, Cape,

between Jakkalswater and Vioolsdrift, 600 m. Sept.

2006, Williamson 6048 (BOL, duplicate at E). [The

specimen SUG 5890 cited by Carter (2002) at STL, now

incorportated into NBG, does not exist. In designating

a neotype, Williamson (2007) cited “'Williamson 6048

(BOL, E)’. Since two specimens are cited, this neotypifi-

cation was invalid and this is rectified here.]

Euphorbia namaquensis was included under E. frie-

drichiae in Bruyns et al. (2006). These two species are

very similar and (among various ‘medusoid’ species

occurring in the arid south of Namibia and north-west-

ern South Africa), they share the feature of particularly

slender branehes which become thieker towards their

bases. The two differ in that the cyathia-bearing pedun-

cles arise in E. friedrichiae at or near the tips of the

branches (the tip of the branch elongating into a pedun-

cle in some cases) around the apex of the plant while

in E. namaquensis the cyathia-bearing peduncles are

shorter (and more densely tuberculate) and arise lower

on the branches mainly in the lower half of the plant.

The cyathia in both are of a similar size but the glands

have longer and more slender processes in E. frie-

drichiae, while the ovary is densely pubescent with short

styles (glabrous to pubescent with often much longer

styles in E. namaquensis). In E. friedrichiae in Namibia,

the capsules often have an unusual array of warts and

slightly raised wing-like ridges along the three angles

while they are quite without these in E. namaquen-

sis. However, these excrescences are usually (though

not always) absent in plants in South Africa of E. frie-

drichiae (from east of Onseepkans), where the capsules

are also often larger than in Namibian plants.

White et al. (1941) expressed doubt as to whether

the two names E. multiramosa and E. namaquensis

represented distinct species. E. multiramosa was also

included under E. fiiedrichiae in Bruyns et al. (2006).

Williamson (2007) made extensive notes on E. multira-

mosa and E. namaquensis. He concluded that they repre-

sented distinet species, since ‘the general appearance of

both plants is quite different.... Cymes in E. multiramosa

are only produced on the leeward aspect mostly from

half to the lower third of the plant.... the cymes are soli-

tary with very short peduncles and the involucral glands

smaller, sessile, horizontally curving outwards with 4-8

marginal processes. The capsules are glabrous and ± 8

mm in diameter. Euphorbia namaquensis has a single or

up to two pairs of cyathia with elongated peduncles at

branch apices and with involueral glands larger, shortly

stipitate, suberect to erect and with 3-6 marginal proc-

esses and capsules densely pubescent ± 10-12 mm in

diameter.’ In practice, the ‘general appearanee’ of plants

from north of Steinkopf (taken to be typical of E. multi-

ramosa) and those from west of Gamoep (taken to rep-

resent E. namaquensis) is identical; all the other features

mentioned are actually very variable within populations.

Consequently E. multiramosa and E. namaquensis differ

only in the glabrous vs. pubescent capsules, though even

this feature has been found to be variable in E. multira-

mosa.

E. namibensis Marloth, Transactions of the Royal

Society of South Africa 1:318 (1909). Type: Namibia,

near Tschaukaib about 31 miles from Angra Pequena,

800 m, Nov. 1908, Marloth 4635 (PRE, holo., K, SAM,

iso.). [The specimen under this number at BOL is from

a different locality (in desert near Ltideritzbucht, 50 m,

Aug. 1909) and so is not part of the same collection,

although it bears the same number.]

E. argillicola Dinter: 27 (1914). Type: Namibia, flats

around Jakkalskuppe, Jan. 1910, Dinter 3145 (SAM,

holo.).

E, namuskluftensis L.C. Leach, The Journal of

South African Botany 49: 189 (1983). Type: Namibia,

Namuskluft, ± 1 200 m, Oct. 1978, Lavranos & Pehle-

mann 20796 (PRE holo.; WIND, iso.).

E. nesemannii R.A.Dyer, Bulletin of Miscellane-

ous Information 1934: 267 (1935). Type; South Africa,

240

Cape, koppie west of Robertson, 300’, Jul. 1930, Nese-

rnami sub Dyer 2441 (GRA, lecto., designated here, K,

isolecto.). [Dyer (1935) cited Dyer 2440 (GRA, K) and

Dyer 2441 (GRA, K), so one has been selected as lecto-

type.]

E. oatesii Rolfe in Oates, Matabeleland Victoria

Falls, ed. 2, appendix V: 408 (1889). Type: Zimbabwe,

Matabeleland, Apr. 1878, F. Oates (K, lecto., designated

here). [Rolfe also cited: Zambia, Rogers 8466 (K); Zim-

babwe, 160 km northeast of Bulawayo, Rand 218 (mis-

sing).]

E. obesa Hook.f, Curtis’ Botanical Magazine 129: t.

7888 (1903). Type: South Africa, Kendrew, near Graaff-

Reinet, 2 000’, Mar. 1897, MacOwan 3153 (K, holo.).

E. symmetrica A.C.W\\\\.q et al.\ 964 (1941). E. obesa

subsp. symmetrica (A. C. White et al.) G.D. Rowley: 97

(1998). Type: South Africa, Cape, 19 miles northwest of

Willowmore on road to Rietbron, Aug. 1939, Dyer 4038

(PRE, lecto., designated here; K, isolecto.). [Dyer did

not specify which of these specimens is the holotype and

so a lectotype is selected.]

E. oxystegia Boiss., Centuria Euphorbiarum: 27

( 1860). Type: South Africa, Cape, between Goedemans-

kraal and Kaus, Drege (S; lecto., designated here; K, P,

W, isolecto.). [Boissier (1860) did not cite a herbarium

here and so a lectotype is selected.]

E. patula Mill., Diet., ed. 8: Euphorbia no. 11 (1768).

Dactydanthes patula (Mill.) Haw., Syn. PI. Succ.: 132

(1812). Medusea patula (Mill.) Klotzsch & Garcke,

Monatsber. Konigl. Akad. Wiss. Berlin 1859: 251

(1859). Neotype (designated here): South Africa, Cape,

sheet 328, specimen (one of two) labelled ‘Grimwood’s

St’ by Haworth (OXF). [There are two specimens on

this sheet in Haworth’s Herbarium at Oxford. The one

selected here is fertile, while the other, labelled ‘My

own’ is sterile.]

E. ornithopus Jacq.: 76, t. 120, fig. 2 (1809). Type:

Jacq., Fragm. Bot., 6: t. 120, figure 2 (1809) (lecto., des-

ignated here).

The name Euphorbia patida Mill, has been a

source of considerable confusion. N.E. Brown (1915:

293) suggested that it was a weak form of E. mauri-

tanica and this was taken up by White et al. (1941:

120), while Carter (2002) referred it to E. tridentata.

Both Brown (1915) and White et al. (1941) consid-

ered, wrongly, that Dactylanthes patula was published

Haworth (1812), while it was merely a new combina-

tion for Miller’s name E. patula. White et al. (1941 ) also

believed that Robert Sweet (1818) described a new spe-

cies ''Euphorbia patula’. However, there he referred to

‘H.S.’, which meant ‘Haworth on Succulent Plants’, i.e.

Haworth (1812). Since this provided a clear reference to

Haworth’s book and hence back to Miller (1768), it did

not constitute publication of a new, and then illegitimate

name Euphorbia patula Sweet, as was assumed in White

et al. (1941) and Carter (2002) but merely referred to

Miller’s E. patida. White et «/.(194l) also considered

that Klotzsch & Garcke (1859) published a new name

Medusea patula but this, too, is wrong and this was also

a new combination for E. patula Mill. Consequently,

Bothalia42,2 (2012)

they missed the fact that Miller’s name E. patida was the

earliest valid name for E. ornithopus.

E. pedemontana L.C. Leach, South African Journal

of Botany 54: 501 (1988). Type: South Africa, Cape,

foot of Matsikamma, Vanrhynsdorp distr., Lavranos &

Bleck 20828 (NBG, holo.).

E. pentagona Haw., The Philosophical Magazine, or

Annals of Chemistry, Mathematics, Astronomy, Natu-

ral History and General Science, Ser. 2, 3: 187 (1828).

Type: South Africa, Cape of Good Hope, received 1823,

Bowie (missing). Neotype (designated here): South

Africa, Cape, Kei River Mouth, Flanagan 2344 (BOL;

duplicates at GRA, PRE). [The painting number 296/926

at K by G. Bond represents a very weak and imper-

fectly developed branch without spines (as noted by

N.E. Brown on the painting) and it is doubtful whether

this is a reasonable lectotype as it could belong to one

of several species. Brown (1915) also doubted whether it

was made from the plant from which Haworth described

the species, since Haworth (1828) mentioned spines and

these are absent from this painting. Rather than use this

painting of somewhat doubtful identity as lectotype, a

neotype has been selected.]

E. pentops Marloth ex A. C. White et al., The Suc-

culent Euphorbieae 2: 963 (1941). Type: South Africa,

Cape, near Komaggas, 10 June 1930, Herre 5562 (PRE,

holo.).

E. pillansii N.E.Br., Bulletin of Miscellaneous Infor-

mation 1913: 122 (1913). Type: South Africa, Cape,

near Doomkloof River, between Muiskraal and Ladi-

smith, Aug. 1907, N.S.Pillans sub BOL 12543 (BOL,

holo.; K, iso.). [N.E. Brown wrote ‘1 piece kept for

Kew’ on the specimen at BOL, which is a much larger

specimen, so this is taken as the holotype.]

E. pillansii var. albovirens A. C. White et al.'. 965

(1941). Type: South Africa, Cape, Paardekop near

Spes Bona, 650 m, 3 Oct. 1925, Marloth 12543 (PRE

0258928-1, lecto., designated here; PRE, isolecto.).

[There are two specimens at PRE, neither was selected

by the authors as type so a lectotype is selected here.]

E. pillansii var. ramosissima A.C. White et al.'. 965

(1941). Type: South Africa, Cape, between Montagu and

Touws River, Aug. 1939, Dyer 4100 (missing).

E. polycephala Marloth, South African Gardening &

Country Life 21: 133 (1931). Type: South Africa, Cape,

near Mortimer, Aug. 1913, Shoesmith sub Marloth 5295

(PRE, lecto., designated here). [Marloth (1931) cited

also Marloth 12644 but this is missing.]

E. polygona Haw., Miscellanea naturalia, sive dis-

sertationes variae ad historiam naturalem spectantes:

184 (1803). Neotype (designated here): South Africa,

Cape, Witpoortsberg, 2 000-3 000’, Aug., Drege 8212

(S 2583; duplicates at BM, HBG-2 sheets, K, MO, P, S,

W-3 sheets). [Haworth (1803) mentioned that E. poly-

gona was described from material introduced before

1790, but nothing was preserved. A neotype has there-

fore been selected.]

Bothalia 42,2 (2012)

241

E. horrida Boiss.: 27 (1860). Type: South Africa,

Cape, Witpoortsberg, 2 000-3 000’, Aug., Drege 8212

(S 2583, lecto., designated here; BM, HBG-2 sheets, K,

MO, P, S, W-3 sheets, isolecto.). [Boissier (1860) did not

cite a herbarium and so a lectotype is chosen.]

E. horrida var. striata A. C. White et al.\ 964 (1941).

Type: South Africa, Cape, 15 miles north of Steytler-

ville, Liickhoff 123 (missing).

E. horrida var. noorsveldensis A.C. White et al.: 965

(1941). Type: South Africa, Cape, 1.3 miles north of

Jansenville, Aug. 1939, Dyer 4010 (PRE, holo.).

E. horrida var. wo/or A.C. White et al.\ 965 (1941).

Type: South Africa, Cape, Kruidfontein, 19 miles from

Willowmore towards Rietbron, Aug. 1939, Dyer 4041

(missing).

E. polygona var. nivea Schnabel: 25 (2011). Type:

South Africa, Long Kloof, Kleinrivier, 508 m, 15 Nov.

2010, Schnabel 1 (GRA, holo.).

E. polygona var. exilis Schnabel: 20 (2012). Type:

South Africa, Eastern Cape, foothills of Kouga Moun-

tains, 65 m, 16 Nov. 2011, Schnabel 4 (GRA, holo.).

E. procumbens Mill., The Gardener’s Dictionary,

ed. 8: Euphorbia no. 12 (1768). Medusea procumbens

(Mill.) Haw.: 134 (1812). Neotype (designated here):

J.Burm., Rar. Afric. PL: t. 10, fig. 1 (1738). [Miller cited

neither material nor figures. The figure designated here

as a neotype was cited by Haworth (1812) under his

‘account’ of Medusea procumbens and so gives an indi-

cation of what was then understood by Miller’s name.

Haworth’s association of this figure with Miller’s name

is unlikely to have been co-incidental. Burmann (1738:

t. 10) referred to the plant in this figure as " Euphorbium

hiimile, procumbens,...' so that it is likely that Miller

adopted Burman’s adjective "procumbens’, as White et

al. (1941) suggested.]

E. pugniformis Boiss.: 92 (1862). Type: J.Burm.,

Rar. Afric. PL: t. 10, fig. 1 (1738) (lecto., designated by

Wijnands 1983).

E. gorgonis A. Berger: 230 (1910). Type: South

Africa, Cape, neither collector nor locality (missing).

[Carter (2002) cited a specimen of Burtt-Davy at PRE

as the type, but this does not exist, nor is there any

evidence that it could possibly be the type of Berger’s

name. In his discussion of E. gorgonis, Berger (1910)

mentioned having obtained plants of the recently

described E. davyi from Burtt-Davy but not that Burtt-

Davy had supplied him with E. gorgonis. This appears to

have been mis-interpreted by Carter (2002).]

The name E. procumbens was not used in Bruyns et

al. (2006). This followed White et al. (1941), who did

not adopt E. procumbens Mill, as the name for these

plants, even though it antedated E. pugniformis (based

on the same figure) by nearly 100 years, apparently

because Miller’s ‘description is too incomplete to permit

of any certainty’ (p. 337) in its identity and ‘that name

cannot be maintained at all’ (p. 338). However, its iden-

tity is clear from Haworth’s references which lead to the

present neotypification and the replacement of E. pug-

niformis by this name. White et al. (1941) assumed that

Sweet’s (1818) use of "Euphorbia procumbens’ was a

new name but, since Sweet (1818) referred to Haworth

(1812) and thus indirectly to Miller, they were not cor-

rect.

E. pseudoglobosa Marloth, South African Garden-

ing & Country Life 19: 191 (1929). Type: South Africa,

Cape, near Krombeks River, Riversdale distr.. Sept.

1933, Muir 4089 (PRE, holo.).

E. frickiana N.E.Br.: 491 (1931). Type: South Africa,

Riversdale div., Ferguson comm. Frick (K, holo.).

E. juglans Compton: 126 (1935). Type: South Africa,

Cape, about 20 miles west of Ladismith, Feb. 1932,

Compton 3951 (BOL, holo.).

E. pseudotuberosa Pax, Bulletin de L’Herbier Bois-

sier, ser. 2, 8: 637 (1908). Type: South Africa, Transvaal,

Pretoria, 1892, Fehr 43 (Z, holo.).

E. pulvinata Marloth, Transactions of the Royal

Society of South Africa 1: 315 (1909). Type: South

Africa, Cape, Queenstown, Marloth 4372 (missing).

Neotype (designated here): South Africa, Cape, Queen-

stown, Nov. 1898, Galpin 2527 (PRE). [The type has

not been located. A neotype from the same locality is

selected here.]

E. quadrata Nel, Jahrbuch der Deutschen Kakteen-

Gesellschaft 1: 42 (1935). Type: South Africa, Cape,

near summit of Stinkfonteinberg, Oct. 1930, Herre sub

SUG 6519 (BOL, holo.). [Carter (2002) cited this speci-

men at STE (now incorporated into NBG) and BOL, but

the former does not exist.]

E. stegmatica Nel: 43 (1935). Type: South Africa,

Cape, Stinkfonteinberg, Oct. 1930, Herre sub SUG 6518

(BOL, holo.). [Although the illustration in Nel (1935)

is clearly of E. oxystegia, the type is a specimen of E.

quadrata.]

E. francescae L.C. Leach: 563 (1984b). Type: South

Africa, Cape, Comellsberg, Sept. 1984, Williamson 3248

(NBG, holo.).

E. restituta N.E.Br., Flora capensis 5(2): 339 (1915).

E. radiata E.Mey. ex Boiss.: 90 (1862), nom. illegit.,

non Thunb. (1800). Type: South Africa, Cape, between

Stinkfontein and Garies, Pillans 5579 (BOL, lecto., des-

ignated here; K, isolecto.). [Brown (1915) also cited:

Schlechter 11098 (BOL); between Zwartdoom R. and

Groen R., Aug, Drege 2941 (missing). The latter is prob-

ably the same as the specimen which Boissier (1862)

cited: between Zwartdoom R. and Groen R., Aug.,

Drege (S, W).]

E. graveolens N.E.Br.: 253 (1915). Type: South

Africa, Cape, between Stinkfontein and Garies, Dec.

1910, Pillans 5579 (BOL 137769, lecto., designated

here; BOL, K, isolecto.). [Brown (1915) also cited:

Bakhuis, Pillans 5486 (K).]

E. schoenlandii Pax, Jahresbericht der Schlesischen

Gesellschaft fur vaterlandische Cultur 82: 24 (1905).

Type: South Africa, Cape, ‘Clanwilliam (Woodifield)’,

242

Bothalia 42,2 (2012)

fl. May 1904, Schonland (GRA, lecto., designated here).

[This specimen was annotated as ‘Co-type’ by Schon-

land and is unlikely to have been seen by Pax. It is pre-

sumed that the other part was sent to Pax and this part

remains missing.]

E. silenifolia (Haw.) Sweet, Hortus Britannicus, ed. 1,

2: 356 (1826). Tithynialus silenifolius Haw.: 61 (1821).

Type: Illustration number 810/147 by T. Duncanson at

K of specimen received 1823 from Cape of Good Hope

collected by Bowie (lecto., designated here).

E. elliptica Thunb.: 86 (1800), nom. illegit., non

Lam (1786). Tithymalus eUipticus (Thunb.) Klotzsch &

Garcke: 69 (1860). Type: South Africa, Cape, Thimberg

(UPS-THUNB 11446, holo.).

Tithymalus hergii Klotzsch & Garcke: 68 (1860).

Type: South Africa, Cape, Bergius (missing).

Tithymalus longipetiolatus Klotzsch & Garcke: 68

(1860). Type: South Africa, Cape, Bergius (missing).

Tithymalus attenuatus Klotzsch & Garcke: 69 (1860).

Type: South Africa, Cape, Bergius (missing).

E. elliptica var. undulata Boiss.: 93 (1862). Neotype

(designated here): Type: Illustration number 810/147

by T. Duncanson at K of specimen received 1823 from

Cape of Good Hope collected by Bowie. [Boissier

( 1 862) cited ‘ Tithymalus silenifolius & Tith. crispus

Haw., revis. pi. Succul. p. 61 {ex descriptionef , so he

took these two names as applying to the same species

and combined them under this variety. This view is not

supported here. By selecting a neotype as above, this

name becomes a synonym of E. silenifolia.^

E. mira L.C. Leach: 10 (1986a). Type: South Africa,

Cape, near Tulbagh, Bayer sub Leach 17175 (NBG,

holo.; K, PRE, iso.).

Although the name E. mira L.C. Leach was main-

tained as a distinct species in Bruyns et al. (2006),

observations of populations of E. silenifolia have made

it clear how this species may begin its growth extremely

early (in February, well before winter) and how narrow

the leaves may be in some populations, often mixed

up with plants with considerably broader leaves. Thus,

while Leach (1986a: 11) believed he had found three,

possibly even four geophytic species of Euphorbia

growing together at the type locality of E. mira, it is

clear from the photograph (Leach 1986a: figure 2) and

the specimens made, that he found E. tuberosa and vari-

ous forms of E. silenifolia at this locality.

E. stellispina Haw., The Philosophical Magazine, or

Annals of Chemistry, Mathematics, Astronomy, Natu-

ral History and General Science, Ser. 2, I: 275 (1827).

Type: Illustration number 803/324 by T. Duncanson at

K of specimen received 1822 from Cape of Good Hope

collected by Bowie (lecto., designated here). [The paint-

ing selected as lectotype was made from the plants seen

by Haworth (of which no material was preserved). There

are two figures of E. stellispina by Duncanson and this

one is selected as the other exhibits very odd growth and

is not representative of the species.]

E. stellispina var. astrispina (N.E.Br.) A. C. White et

al.: 716 (1941). E. astrispina N.E.Br.: 355 (1915). Type:

South Africa, Beaufort West distr., Willowmore side,

Brauns 1711 (K, holo.).

E. susannae Marloth, South African Gardening &

Country Life 19: 191 (1929). Type: South Africa, Cape,

Phisantefontein, Oct. 1923, Muir 2762 (BOL 137790,

lecto., designated here; BOL, PRE, isolecto.). [Marloth

(1929) cited also: Marloth 12155 (NBG, PRE).]

E. systyloides Pax subsp. porcaticapsa S. Carter,

Kew Bulletin 45: 336 (1990). Type: Zimbabwe,

Hurungwe distr., Zambesi Valley, Rifa R., 520 m, 24

Feb. 1953, Wild 4085 (K, holo.; EA, SRGH, iso.).

E. trichadenia Pax, Botanische Jahrbiicher fur Sys-

tematik 19: 125 (1894). Type: Angola, Lunda, between

Kimbundo and the Quango, Sept. 1876, Pogge 116

(missing). Neotype (designated here): Angola, Huilla,

near Lopollo towards Nene, Oct.-Nov. 1859, Welwitsch

282 (BM; duplicates at G, K). [Although Carter (2002)

cited this specimen as being at B, there is no such mate-

rial there.]

E. benguelensis Pax: 741 (1898). Type: Angola,

Huilla, source of Luala, Antunes 362 (missing).

E. subfalcata Hiem: 948 (1900). Type: Angola,

Huilla, near Lopollo towards Nene, Oct.-Nov. 1859,

Welwitsch 282 (BM, holo.; G, K, iso.).

E. gossweileri Pax: 88 (1909). Type: Angola, Maland-

sche, Gossweiler 994 (K, lecto., designated here). [There

is no sign that Pax saw this specimen although N.E.

Brown wrote ‘Type’ on it. Consequently it is selected as

lectotype.]

E. trichadenia var. gibbsiae N.E.Br.: 524 (1911-12).

Type: Zimbabwe, near Isotye, Matopos, 5 000’, Feb.

1905, Gibbs 234 (BM, holo.; K, iso.). [Brown (1911-12)

cited: Gibbs 234 (BM, K) and Victoria, Munro. Two col-

lections of Munro have been located, namely Munro 141

(BM) and Munro 1467 (BM). However, he wrote ‘Type’

on Gibbs 234 (BM) and ‘From the type’ on Gibbs 234

(K) and nothing of this kind on the Munro collections

so it is clear that the Gibbs specimens are holotype and

isotype respectively.]

E. tridentata Lam., Encyclopedic methodique

2(2): 416 (1788). Medusea tridentata (Lam.) Klotzsch

& Garcke: 251 (1859). Type: South Africa, collector

unknown (P-LAM P00381880, holo.; K, iso.).

E. anacantha Aiton: 136 (1789). Dactylanthes

anacantha (Aiton) Haw., Syn. PI. Succ.: 132 (1812).

Type: Illustration in J. Burm., Rar. Afric. PI.: t. 7, fig.

2 (1738) (lecto., designated here). [This figure and one

by D’Isnard (1720) were cited by Aiton (1789), The one

selected as lectotype here corresponds more closely to

the concept of E. tridentata adopted here, while that of

D’Isnard is somewhat more suggestive of E. patula.]

E. tuberosa L., Species Plantarum I: 456 (1753).

Tithymalus tuherosus (L.) J. Hill, Hort. Kew.: 172/3

(1768). Type: Illustration in J. Burm., Rar. Afric. PI.: t.

4 (1738) (lecto., designated here). [This figure was cited

by Linneaus ( 1 753). It was also cited by Carter (2002) as

‘T: icono’ but this does not constitute valid lectotypifica-

tion.]

Bothalia 42,2 (2012)

243

E. crispa (Haw.) Sweet: 356 (1826). Tithymaliis cris-

pus Haw.: 61 (1821). Type: none located.

E. tugelensis N.E.Br., Flora capensis 5(2): 335

(1915). Type: South Africa, Natal, near Tugela River,

received July 1865, Gerrard 1626 (K, holo.; W, iso.).

E. wilmaniae Marloth, South African Gardening &

Country Life 21: 133 (1931). Type: South Africa, Cape,

Boetsap, Pagan sub Marloth 6125a (PRE, lecto., des-

ignated here). [Marloth (1931) cited two specimens:

Boetsap, MacGregor Museum 2337 (missing); Lekker-

sing, Marloth 12441 (PRE). The latter belongs to E.

celata (Leach 1984a, Bruyns 1992). A specimen anno-

tated exactly as the first has not been found but Wilman

(1946) cited this collection as ‘Boetsap, 2337 Pagan’

and so the specimen ‘Boetsap, Pagan sub Marloth

6125a (PRE)’ is strongly suspected to be this collection

and is thus taken as the lectotype.]

E. planiceps Marloth ex A. C. White et al.\ 963 (1941 ).

Type: South Africa, Cape, farm near Griquatown, Sept.

1939, Venter (BOL, lecto., designated here). [White et

al. (1941) designated the collection by Venter as ‘type’

and that by Mrs Cooke (missing) as ‘type of inflores-

cence’ so a lectotype is designated. The name was first

used by Marloth (Wilman 1946).]

Excluded Names

E. aggregata A. Berger, Sukk. Euph.: 92 (1906a).

Type: South Africa. Cape (missing). [No preserved

material has been found of this species and it is difficult

to be sure whether it falls under E. ferox or E. pulvinata

or refers to the intermediates between them that occur

widely over the eastern Karoo.]

E. curvirama R. A. Dyer, Rec. Albany Mus, 4: 104

(1931). Type: South Africa, Cape, 28-30 miles from

Grahamstown towards Peddie, Apr. 1928, Dyer 1403

(PRE, holo.; GRA, K, iso.). [This is considered to be a

hybrid, possibly between E. caerulescens and E. trian-

gularis.]

E. inconstantia R.A.Dyer, Rec. Albany Mus. 4: 93

(1931). Syntypes: Hellspoort, Oct. 1928, Dyer 1076

(GRA); Grahamstown, Aug. 1927, Dyer 1076 (GRA);

10 miles from Grahamstown on Queen’s road, Nov.

1926, Dyer 669 (GRA); Oct. 1927, Dyer 1077 (GRA);

Nov. 1926, Dyer 669a (GRA). [This is considered to be

a hybid, possibly between E. heptagona and E. poly-

gona.]

E. mamillosa Lem., Illustr. Hort. II, misc.: 69 (1855).

Type: unknown. [Lemaire (1855) listed dnamillosa

Nob.’, of unknown origin, among 18 names in Sect.

Aculeatea and provided a Latin diagnosis for it. White

et al. (1941) listed the name as a synonym of E. squar-

rosa (= E. stellata), but it is hard to justify this from the

details that Lemaire gave. White et al. (1941) also listed

the name " Anthacantha mamillosa Lem.’ and gave the

same location as its place of publication, but this name

does not exist.]

E. multifida FI. cap. 5(2): 253 (1915). Type:

South Africa, Natal?, 1905, Anon sub 10483 (NH,

holo.). [The type of£. multifida consists of several inflo-

rescences only, is of unknown origin (though suspected

of coming from ‘Natal’) and the collector is unknown.

It is not, at present, identifiable with certainty with any

known species and so is placed among the excluded

names.]

E. parx’imamma Boiss. in DC., Prodr. 15(2): 86

(1862). [Boissier (1862) cited no material other than a

sterile plant apparently in cultivation under the name E.

caput-medusae, which may have originated at the Cape

of Good Hope. The description is meagre and identifica-

tion remains uncertain.]

E. scolopendria Donn, Hort. Cantab., ed. 3: 88

(1804). [Donn mentioned only that this was ‘flat-leaved’

and flowered Jun.-Aug. No region of origin was given.]

E. viminalis N.L.Burm., Prodr. FI. Cap.: 14 (1768).

[Both White et al. (1941) and Boissier (1862) cite this

name. Actually no description or diagnosis was given

by Burman and he merely listed Euphorbia viminalis

of Linneaus, which is the basionym of Sarcostemma

viminale (L.) R.Br. (Apocynaceae). Here Burman

(1768) cited ‘Alp. aegypt. t. 190. Dill elth. t. 368’ and

he appears to have copied these references directly from

among the five given by Linneaus (1753) for E. vimi-

nalis L. (= Sarcostemma viminale). In fact these refer-

ences are wrong. In Alpini (1735) there is no t. 190, but

the figure referred to is t. 53 on page 190. This figure is

the lectotype of S. viminale, selected by Liede & Meve

(1993), though it is wrongly cited there too. Dillen’s

Hortus Elthamensis (Dillen 1732) had only 324 plates

in it and here page 386 was meant, where there is no

plate. This was again cited incorrectly in Liede & Meve

(1993).]

E. viperina A. Berger, Monatsschr. Kakteenk. 12: 39

(1902a). Type: South Africa, Cape of Good Hope?, col-

lector unknown (missing). [White et al. (1941) placed

E. viperina under E. inermis. However, the description

of Berger does not correspond closely to what we know

today as E. inermis. No type has been located for E.

viperina. Berger (1902a) compared the inflorescences of

E. viperina to those of E. caput-medusae and E. parvi-

mamma, but in fact the inflorescences of the latter were

never described and it is not clear that what he called E.

parvimamma (Berger 1899) corresponds to Boissier’s

concept of it.]

ACKNOWLEDGEMENTS

The curators of the herbaria B, BM, BOL, G, GRA,

K, KMG, M, NBG, NY, OXF, P, PRE, S, SBT, SAM, W,

WIND, WU and Z are thanked for access to the material

in their care. Christiane Anderson, University of Michi-

gan is thanked for copies of many pieces of little-known

literature and for much assistance with, and helpful dis-

cussion of, nomenclatural matters concerning several

of the names in Euphorbia. Paul E. Berry, University

of Michigan, with funds from the U.S. National Sci-

ence Foundation PBI program (award # DEB-0616533),

assisted with the costs of a trip to examine specimens in

some European herbaria during which many types were

located. He also provided invaluable advice on the sta-

tus of many other types, among extensive comments on

244

Bothalia 42,2 (2012)

an earlier draft of this paper. S.P.Bester, SANBI, Preto-

ria, helped to elear up certain problems in the collecting

books of R.A. Dyer and Marloth. Gill Challen, Royal

Botanic Gardens, Kew provided assistance with locating

illustrations and many relevant specimens in K. Fiona

Jones, Jagger Library, University of Cape Town, helped

trace certain works online. Serena Mamer, Oxford Uni-

versity, checked on specimens attributed to Haworth.

This research was also partially funded by grants from

the National Research Foundation and from the Uni-

versity Research Committee of the University of Cape

Town.

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Bothalia 42,2: 247-250 (2012)

FSA contributions 20: Asteraceae: Anthemideae: Inezia

N. SWELANKOMO*

INTRODUCTION

The genus Inezia E. Phillips comprises two species

endemic to eastern southern Africa, namely /. intergri-

folia (Klatt) E. Phillips from Mpumalanga and Swaziland

and I. speciosa Brusse from Limpopo. The type species

I. integrifolia was originally described by Klatt (1896)

in the genus Lidbeckia but Philips (1932) later accom-

modated it within a new monotypic genus Inezia (Table

1 ). A second species, I. speciosa was later described by

Brusse (1989). The genus is placed within the subtribe

Cotulinae of tribe Anthemideae (Himmelrich et al. 2008;

Oberprieler et al. 2009). Inezia is distinguished from

Adenanthelliim and HiUiardia by having ovate or pinna-

tisect leaves and oblong, 4-angled cypselae with bicellu-

lar glands scattered on the outer surface. In the previous

subtribal classification of Bremmer (1994), Inezia was

placed within the now sub-assumed subtribe Thamino-

phyllinae, together with Adenanthelliim B.Nord., Lid-

beckia P.J.Bergius and Thaminophyllum Harv. Nordens-

tam (1976, 1987), when describing the genus HiUiardia

B.Nord., suggested that it had affinities to Inezia and

Adenanthelliim on the basis of the similar ray florets

with bifid or emarginate limbs and the often reduced

perianth tube with a mix of eglandular hairs and bicel-

lular glands on the margins, the distinctive branched

venation and minutely papillate upper surface of the

perianth limbs, and the linear, flattened, epappose cypse-

lae with ciliate margins and scattered bicellular glands

on the outer surface (Figure lA-D). This relationship

is supported by phylogenetic analyses of cpDNA ndhF

sequence data (Elimmelrich et al. 2008). Analyses of

nrlTS sequence data, however, suggests a close relation-

ship between Inezia and HiUiardia, but places Thami-

nophylhim rather with Lidbeckia. A close relationship

between Lidbeckia and Thaminophyllum was also sug-

gested by Bremer & Humphries (1993) based on the

pilose receptacles and myxogenic cells on the cypselae.

TABLE 1. — Differences between Lidbeckia and Inezia.

■ National Herbarium, Biosystematics Research and Biodiversity Col-

lections Division, South African National Biodiversity Institute, Private

Bag XI 01, Pretoria, 0001. E-mail: n.swelankomo @sanbi.org.za.

MS. received: 201 1-06-30.

In this paper a detailed taxonomic treatment of the

two species of Inezia is presented, including a key to the

species, typification, distribution maps and illustrations.

MATERIAL AND METHODS

All the Inezia, HiUiardia, Adenanthelliim, Thamino-

phyllum, and Lidbeckia specimens at PRE were studied.

Digital images of type collections were, where neces-

sary, accessed on the Aluka digital library (http://plants.

jstor.org, accessed July 2011). HiUiardia ziiiirbergensis,

I. integrifolia, I. speciosa, Lidbeckia pectinata, and Lid-

beckia qiiinqiieloba specimens were examined by means

of the scanning electron microscope (SEM). All samples

were dry and were not chemically treated before being

sputter-coated with gold-palladium.

MICROMORPHOLOGY

Inezia is characterised by the presence of large sessile

glands on the ovary, cypselae, perianth tube and perianth

lobes of both ray and disc florets. SEM study reveals

that the large sessile glands are bicellular. The epidermal

cells on the upper surface of the rays of I. integrifolia

are round and radially striate with thick walls (Figure

lA-D).

TAXONOMY

Inezia E. Phillips in Kew Bulletin 6: 297 (1932);

Bremer & Humphries: 146 (1993); Bremer: 471 (1994);

Herman et a/.: 145 (2000). Type species: I. integrifolia

(Klatt) E. Phillips.

Subshrubs with woody rootstock, stems branching

distally, terete, sparsely pilose to villous. Leaves alter-

nate, entire or pinnatisect, glandular-pilose. Capitiila

solitary, terminal, radiate, pedunculate. Involucre cam-

panulate; involiicral bracts in 3 or 4 rows, imbricate, ±

linear, pilose, inner bracts with membranous tip; recep-

tacle conical, epaleate. Ray florets female-fertile or

sterile, yellow or white, with bicellular glands scattered

over the surface; corolla tube short or absent with mix

of eglandular hairs and bicellular glands on the mar-

gins, limb obovate, 2-toothed, three to four x longer

than tube. Style branched, truncate, with stigmatic areas

lateral; ovary with bicellular glands. Cypselae linear,

flattened, margins ciliate and with scattered bicellular

glands on the outer surface, non-myxogenic; pappus

absent. Disc florets bisexual, fertile, yellow with bicel-

lular glands scattered over the surface, narrowly funnel-

shaped, corolla tube somewhat compressed, winged,

4-lobed, two lateral lobes ± cucullate, dorsal and ventral

lobes ovate. Stamens 4, filament collars with 6-8 rows

of cells; anthers minutely caudate, with ovate to oblong

apical appendages; style terete, somewhat swollen at

248

Bothalia 42,2 (2012)

FIGURE I — A, B, ray floret of I.

integrifolia, Taylor 1982: A,

surface with minute papillae

and bicellular glands; B, sur-

face at higher magnification;

C, ray florets of Lidbeckia

pectinata, Esterhuysen 23088

showing radially striations;

D, Hutchinson 2423: base of

perianth limb and cypselae

showing mix of eglandular

hairs and bicellular glands; E,

disc floret of Lidbeckia pecti-

nata, Esterhuysen 23088: E,

surface with bicellular glands.

base, branches oblong, truncate, laterally stigmatic; cells

of stylopodium thick walled; ovaty four ribbed, flat-

tened, bicellular gland scattered throughout. Cypselae

oblong, 4-angled, flattened when matured, with bicel-

lular glands scattered on the outer surface, non-myxo-

genic; pappus absent.

2 spp., northeastern South Africa and Swaziland

Etymology, the genus name Itiezia is named in hon-

our of Ms Inez Clare Verdoorn ( 1896-1989), appointed

to the Division of Botany and Plant Pathology as herbar-

ium assistant in 1917 and later in charge of the National

Herbarium as Senior Professional Officer (1944-1951).

She is commemorated also in Aloe verdoorniae Rey-

nolds, Seuecio verdoorniae R.A.Dyer, Teclea verdoorn-

iana Exell & Mendon^a (Glen & Germishuizen 2010).

Key to species

1. All leaves simple; peduncles up to 50 mm long; ray florets

yellow; Mpumalanga and Swaziland I . /. integrifolia

1.’ Lower leaves pinnatisect, upper leaves simple; peduncles

more than 200 mm long; ray florets white; Limpopo

2.1. speciosa

1. I. integrifolia (Klatt) E. Phillips in K.ew Bulletin

6: 297 (1932). Lidbeckia integrifolia Klatt: 840 (1896).

Type: [Mpumalanga], Saddleback Mountain, 4 000’

[1 312 m], Dec. 1 890, Galpin 1174, (PRE, lecto.!, des-

ignated here; K — Aluka image!, SAMI, isolecto.). [The

Bothalia 42,2 (2012)

249

collection by Galpin is chosen here from the three col-

lections cited by Klatt in the protologue as it has more

precise collection details.]

Subshrub from woody rootstock, 0.2-0. 6 m high,

stems branching distally, sparsely pilose to villous.

Leaves alternate, sessile, ovate, 6.5-22.0 x 3. 0-5. 5

mm, entire, acuminate, dull green, moderately to

densely pilose. Capitiila solitary, pedunculate, pedun-

cle 20-50 mm long, ribbed, glandular-pilose; involucre

campanulate, 5-12 mm diam.; invohicral bracts in 2

or 3 series, imbricate, sparsely to densely hairy, grey-

ish with red-purple tips or rimmed at top with purple;

outer bracts linear, 3. 5^.0 x i.Q mm, moderately to

densely pilose, inner bracts oblong with membranous

tips, 4-7 X 0.5-1. 2 mm; receptacle epaleate. Ray flo-

rets female-fertile, scarcely longer than involucre, yel-

low with bicellular glands scattered over the surface;

corolla tube 1.5 mm long, base of perianth with mix of

eglandular hairs and bicellular glands on the margins,

limb up to 4.5 mm long, 2-lobed, adaxial epidermal cells

round with thick walls and radial striations; style 1. 0-1.1

mm long, branches 0.1 mm long. Cypsela 1.0-1. 2 mm

long, ciliate along margins, with scattered bicellular

glands on the outer suface. Disc florets bisexual, fer-

tile, yellow, 1. 5-2.0 mm long, bicellular glands scat-

tered over the surface, lobes 0.1 mm long; style 1. 8-2.0

mm long, branches 0.18-0.20 mm long; ovaiy 1.0 mm

long. Cypsela 1.2-1. 5 mm long, 4-angled and laterally

flattened, ciliate along margins, with bicellular glands

scattered on the outer surface, non-myxogenic; pappus

absent. Flowering time: Nov.-Apr. (Figures lA-E).

Distribution and habitat'. Inezia intergr {folia occurs in

Mpumalanga, from around Piet Retief and Komatipoort,

north to Pilgrim’s Rest, and around Mbabane in Swazi-

land. from 650-2 152 m (Figure 2). The species favours

montane grasslands, especially humus-rich soils.

Notes: Inezia intergrifolia is distinguished from I.

speciosa by the consistently simple leaves along the

entire length of the branch, the shorter peduncles up to

50 mm long, and the smaller involucre 5-12 mm long

FIGURE 2. — Distribution of Inezia inlegrifolia, ■; and /. speciosa, •.

with red or purple tipped bracts arranged in 2- or 3 rows,

and the yellow ray florets.

Conservation status: LC (Least Concern) (Raimondo

et al. 2009).

Additional specimens

MPUMALANGA. — 2430 (Pilgrim’s Rest): Mount Sheba, (-DC),

14 Feb. 1982, Brenan 14962 (PRE); Ohrigstad Dam Nature Reserve,

(-DC), 16 Feb. 1972, Jacobsen 2321 (PRE); Pilgrim's Rest on hill-

sides, (-DD), 12 Mar. 1937, Galpin 14420 (PRE); Graskop, (-DD),

18 Jan. 1921, b-\’ine & Irvine 11421 (PRE); Jan. 1920, Rogers 23592

(PRE). 2530 (Lydenburg): Mount Anderson, summit peak, (-BA),

9 Mar. 1933, Galpin 13757 (PRE); Mount Anderson, (-BA), 16 Jan.

1952, Prosser 1800 (PRE); Vertroosting Nature Reserve, 12 km S of

Sabie, (-BB), 25 Feb. 1972, Muller 2469 (PRE); Nelspruit, Wonder-

kloof Nature Reserve, (-BC), 18 Nov. 1974, Elan-Puttick 183 (PRE);

Witklip, (-BD), 17 Jan. 1974, Kluge 444 (PRE); Nelspruit, slopes of

Amajuba Mountain (Schagen), (-BD), Dec. 1934, Liehenberg 3320

(PRE); Kaapsehoop, Devil’s Kantoor, (-DB), 1 1 Jan. 1924, Pole Evans

998, (PRE); Barberton, Nelshoogte Forest Station, (-DD), 8 Dec.

1953, Codd 8132 (PRE); Barberton, Thomcroft Nature Reserve, (-

DD), 6 Jan. 1972, Muller 2274 (PRE); Nelsberg, (-DD), 26 Feb. 1936,

Taylor 1982 (PRE). 2531 (Komatipoort): Sheba Mine, Colombo 365

JU Farm 15 KM. S of Sheba Mine, (-CA), 1980, Fourie 230 (PRE);

mountain top between Louw’s Creek and Maid of the Mist, (-CA), 5

Jan. 1929, Hutchinson 2423 (PRE); Malelane, (-CB), Nov. 1924, Mur-

phy 106, (PRE); 10 miles [16,1 km] SE Barberton on road to Have-

lock. (-CC), 9 Dec. 1953, Codd 8165 (PRE); Barberton, Oosterbeek

Farm, 5 km S Barberton, (-CC), 15 Nov. 1977, De Souza 628 (PRE);

on Barberton-Havelock road, 14 miles [22.4] from Barberton, 9 Feb.

1962, Ihlenfeldt 2432 (PRE); near top of mountain behind Barber-

ton village, (-CC), 18 Feb. 1931, Liebenberg 2420 (PRE); Ida Doyer

Nature Reserve, 38 km SE Barberton, (-CC), 7 Dec. 1971, Muller

2050 (PRE); Barberton, (-CC), Jan. 1907, Thomcroft TRV4973

(PRE); Barberton, Tinie Louw Nature Reserve, (-CC), 15 Jan. 1983,

Venter 9176 (PRE); W of Ngodwana, Hemlock Tulloch Mhor Nature

Reserve, (-DA), 15 Feb. 2003, McMurtiy 11157 (PRE). 2730 (Piet

Retief): Piet Retief, 15 km from Piet Retief on road to Amsterdam, (-

BB), Mar. \913, Arnold 245 [VRE).

SWAZILAND.— 2631 (Mbabane): Ngwenya Hills, (-AA), 30 Jan.

1957, Compton 26527 (PRE); Malolotja Nature Reserve, above Forbes

Reef Dam, old kraal, (-AA), 2 Dec. 1986, Heath 533 (PRE); Endin-

geni. Middle veld, (-AB), 18 Dec. 1970, Barrett 551 (PRE); Hhohho,

Malandzela area; road to Maphalaleni, 14 km from Nyokane (tar road)

above Komati River and below Enkaba Trig beacon, (-AB), 25 Jan.

1994, Hobson 20 75 (PRE); Mbabane, (-AC), Dec. 1905, Bolus 12012

(PRE); Black Umbuluzi Valley, 9 km from Mbabane to Balegane near

mission falls, (-AC), 27 Dec. 1985, Brusse 4344 (PRE); Mbabane,

(-AC), Jan. 1905, Burtt Davy 2868 (PRE); 17 Jan. 1951, Compton

22463 (PRE); Mbabane, Ukutula, (-AC), 6 Jan. 1956, Compton 25291

(PRE); Bomvu Ridge, (-AC), 5 Jan. 1962, Compton 31193 (PRE);

Mbabane Area, (-AC), 7 Dec. 1960, Dlamini PH31853 (PRE); Usutu

Forest, (-CA), Dec. 1974, Watson 8 (PRE).

2. I. speciosa Brusse in Bothalia 19: 27-29 (1989).

Type: [Limpopo], Iron Crown Mountain near Haenerts-

burg, 5 500’ [1 804 m], LE.Codd 9440, 24 Jan. 1956

(PRE, holo!; K [2 sheets] — Aluka image!, MO — Aluke

image!, iso.).

Subshrub up to 0.45 m., main stem simple to

2-branched at base but well branched above, erect, hir-

sute, sparsely leafy, branches simple, ascending densely

leafy, hirsute. Leaves dimorphic, lower impari-pinnati-

sect, upper linear-lanceolate, up to 20 mm long, moder-

ately to densely hirsute, margins involute. Capitiila soli-

tary, radiate, pedunculate, peduncle 225-290 mm long,

ribbed, hirsute; involucre campanulate, 20-25 mm long

when pressed; invohicral bracts in 3 or 4 series, imbri-

cate, narrowly lanceolate, brownish-red, hirsute on outer

surface, glabrous on inner surface, outer 5. 0-6.0 xi.Q-

1.5 mm, inner 6. 0-9. Ox 1.0- 1.5 mm. Ray florets 25-30,

250

Bothalia42,2 (2012)

female or, white with bicellular glands scattered over the

surface, corolla tube 1.0 mm long, limb 3-lobed base of

perianth limb with mix of eglandular hairs and bicellular

glands on the margins, adaxial epidermal cells obtusely

acuminate, radially slightly striate; ovary not well

developed, with scattered bicellular glands. Disc florets

bisexual, fertile, yellow, 2. 5-3.0 mm long, with scattered

bicellular glands on the outer surface, lobes 0.4-0. 5 mm

long; style 2.5 mm long, branches 0.6 mm long; ovary

1.0-1. 5 mm long. Cypselae 1.5-2. 5 mm long, 4-angled,

laterally flattened, ciliate along margins with scattered

bicellular glands on the outer surface, non-myxogenic;

pappus absent. Flowering time; Nov.-Jan.

Distribution and ecology: Inezia speciosa is known

only from the type locality near the summit of Iron

Crown Mountain near Haenertsburg, 1 804 m (Figure 1),

where it has been collected on two occasions. It favours

quartzite rock, especially grassy slopes which are rich in

forb species with scattered trees and shrubs (Mucina &

Rutherford 2006).

Notes: Inezia speciosa can be distinguished from

I. integrifolia by its dimorphic foliage, with the lower

leaves pinnatisect and the upper simple, the longer

peduncles more than 200 mm long, the larger involucre

20-25 mm long with bracts in 3or 4 series, and the white

ray florets.

Conservation status: EN Blab (Hi) + 2ab (iii) (endan-

gered) (Raimondo et al. 2009).

Additional specimen examined

LIMPOPO. — 2329 (Polokwane): Wolkberg, Iron Crown, just W of

summit, (-DD), 29 Nov. 1980, McMurtry 4208 (PRE).

ACKNOWLEDGEMENTS

My colleagues from the South African National Bio-

diversity Institute: R Herman for guidance and discus-

sions and Hester Steyn for producing the distribution

map. Referees are thanked for their valuable comments.

The curator and staff of Harvard University Herbarium

and SAM are also thanked for sending digital images of

type material.

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VELDTMAN, R., LADO, T.F., BOTES, A., PROCHES, S., TIMM,

A.E., GEERTSEMA, H. & CHOWN, S.L. 2011. Creating novel

food webs on introduced Australian acacias: indirect effects of

galling biological control agents. Diversitv and Distributions 17:

958-967.

258

Bothalia 42,2 (2012)

VERSTRAETE, B„ VAN ELST, D., STEYN, H., VAN WYK, A.E.,

LEMAIRE, B„ SMETS, E. & DESSEIN, S. 2011. Endophytic

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WILLOUGHBY, S. 2011. SANBI participates in Impact Assessment

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sanbi-participates-impact-assessment-conference-mexico.

WILSON, J.R.U., GAIRIFO, C., GIBSON, M.R., ARIANOUTSOU,

M., BAKAR, B.B., BARET, S., CELESTI-GRAPOW, L.,

DITOMASO, J.M., DUFOUR-DROR, J.M., KUEFFER, C.,

KULL, C.A., HOFFMANN, J.H., IMPSON, F.A.C., LOOPE,

L.L., MARCHANTE, E., MARCHANTE, H., MOORE, J.L.,

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ical control: global efforts to limit Australian acacia invasions.

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WITBOOl, H. 201 1-01 . Aloe zebrina Baker (Asphodelaceae). Internet 3

pp. http://www.plantzaffica.com/plantab/aloezebrina.htm.

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net 3 pp. http://www.plantzafrica.com/plantqrs/strelitziacaudata.

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(Strelitziaceae). Internet 3 pp. http://www.plantzaffica.com/

plantqrs/strelitziareginae.htm.

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strelitzjun.htm.

Bothalia 42,2: 259-269 (2012)

Guide for authors to Bothalia

This guide is updated when necessary and includes an

index. Important points and latest additions appear

in bold type.

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 South Afri-

can National Biodiversity Institute (SANBl), 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 con-

tents, authors and subjects are published annually.

1 EDITORIAL POLICY

1.1 Bothalia welcomes original papers dealing with

flora and vegetation of southern Africa and related sub-

jects. Full-length papers and short notes, as well as

book reviews and obituaries of botanists, are accepted.

The editor should be notified that an article is part of a

series of manuscripts; please submit a list of the parts of

a series; all parts should preferably be published in one

journal.

1.2 Submission of a manuscript to Bothalia implies

that it has not been published previously and is not being

considered for publication elsewhere.

1.3 Authors whose first language is not English are

requested to have their MS edited by an English speaker

before submission.

1.4 Articles are assessed by referees, both local and

overseas. Authors are welcome to suggest possible ref-

erees to judge their work. Authors are responsible for the

factual correctness of their contributions. Bothalia main-

tains an editorial board (see title page) to ensure that

international standards are upheld.

1.5 Page charges: as stated in our notification

included in volume 23,1 (May 1993), MSS submit-

ted for publication in Bothalia are subject to payment

of page charges of R 125,00 per printed page, VAT

included. The following are exempt from these charges:

1, SANBl members; 2, persons/institutions who have

been granted exemption by the Executive Committee

of the SANBl; 3, authors of contributions requested by

the Editor; 4, contributors to the column "FSA contribu-

tions’. The Editor’s decision on the number of pages is

final. An invoice will be sent to the author, who must

arrange for payment as soon as possible to SANBl, Pub-

lications Section, Private Bag XI 01, Pretoria 0001.

1.6 Deadline dates for submission of MS: for possi-

ble inclusion of the MS for the May issue — May of the

previous year, and for the October issue — October of the

previous year.

2 REQUIREMENTS FOR A MANUSCRIPT

2.1 Only electronic submission of manuscripts will

be accepted. Manuscripts do not need to be provided

in hard copy, and manuscripts that are submitted in

hard copy only will not be processed further until the

electronic version is received by the Production Editor.

Manuscripts should be e-mailed to the Production Edi-

tor (y.steenkamp(^sanbi.org.za), along with images of

figures of sufficient quality to use for refereeing pur-

poses (but small enough to e-mail to referees, generally

less than 2 MB per image). Higher quality images, if

too large to e-mail, should be provided on a CD, hand-

delivered, couriered, or mailed to the Production Editor

(see 23. Address of Production Editor, below) for pub-

lication purposes. Line drawings may be provided on a

CD (scanned in at 1200 dpi, in bitmap TIE format) or

originals provided to the Publications Section of SANBl

to assist with the scanning (see 12.3 below for more

details).

2.2 Papers should confonn to the general style and

layout of recent issues of Bothalia (from volume 26

onwards).

2.3 Material should be presented in the follow-

ing sequence: title, name(s) of author(s), address(es) of

author! s) and mention of granting agencies, keywords

and abstract.

2.4 The sequence continues with Introduction and

aims. Contents (see 8), Material and methods. Results,

Interpretation (Discussion), Specimens examined (in

revisions and monographs). Acknowledgements, Refer-

ences, Index of names (recommended for revisions deal-

ing with more than about 15 species). Tables, Captions

of figures and figures. In the case of short notes, obitu-

aries and book reviews, keywords and an abstract are

omitted.

2.5 All pages must be numbered i.e. typed consecu-

tively on the top right-hand comer of the page, begin-

ning with the first page to those with references, tables,

captions of figures and figures.

2.6 Special characters: use your own word or code

that is unique and self-explanatory, enclosed between

angle brackets, e.g. <mu>m for pm. Please supply us

with a list of the codes.

2.7 Use a non-breaking space (in MS Word — Ctrl,

shift, space) to keep two elements together on the same

line, e.g. 3 500.

2.8 Do not justify lines.

2.9 Do not break words, except hyphenated words.

2.10 A hyphen is designated as one dash, with no

space between the letter and the dash, e.g. ovate-lanceo-

late. See also 17.7.

2.11 An N-dash is typed in MS Word code (alt -i-

0150) or as three hyphens with no space between the

letter and the hyphen, e.g. 2 5 mm (typeset, it looks

like this, 2-5 mm). See also 17.7.

2.12 An M-dash is typed in MS Word code (alt +

0151) or as two hyphens with no space between the let-

ter and the hyphen, e.g. computers- -what a blessing!

(typeset, it looks like this: computers — what). See also

17.7.

2.13 Do not use a double space anywhere between

words, after commas, full stops, colons, semicolons or

exclamation marks.

260

Bothalia 42,2 (2012)

2.14 Use lower case x as times sign, with one space on

either side of the x, e.g. 2x3 mm.

2.15 Use single (not double) opening and closing

quotes, e.g. the so-called ‘stiffy’ refers to a rigid diskette.

In MS Word the codes are alt + 0145 and alt + 0146.

2.16 Keys — put only three leader dots before number

of taxon (with one space before and after each dot),

regardless of how far or near the word is from the right

margin, e.g. ... R. ovata (see 13.18).

3 REQUIREMENTS FOR ELECTRONIC FILES

3.1 USE NORMAL STYLE ONLY.

3.2 Provide electronic files on CD or send by e-mail

to y.steenkamp(^sanbi.org.za.

3.3 Data should be in MS Word and in NORMAL

style throughout.

3.4 All lines, headings, keys, etc., should start flush

at the margin, therefore no indentations, footnotes,

tabs or styles of any kind.

3.5 In MS Word, italics and bold should be used

where necessary.

3.6 Paragraphs and headings are delineated by a car-

riage return <ENTER> but no indentation.

3.7 Graphics i.e. drawings, graphs or photographs:

submit in a separate file, do not include it in the text.

3.8 Image files with a bigger file size than 4MB can-

not be e-mailed as the SANBI has a 4MB limitation on

the network’s firewall at Head Office. Piles smaller than

4MB should be e-mailed to: y.steenkamp@sanbi.org.za.

Image files bigger than 4 MB should be provided on a

CD or can be copied to the SANBI FTP site by using the

following link: ftp://ftp.sanbi.org/incoming/. Permission

needs to be obtained to copy material to the FTP site.

Please contact the Production Editor for details.

3.9 Provide an image file originated in Corel Draw

(version 14 or lower), as a CDR file, with fonts con-

verted to curves. Submit image files originated in other

drawing programmes as encapsulated postscript files

(EPS). The conversion to TIE or other file extensions

will be accommodated by SANBI Graphics (see 12.2-

12.4).

3.10 If extensive changes to image files are proposed

by the editor, the author will be contacted and the spe-

cific image file will have to be re-submitted after the

indicated corrections have been implemented.

3.11 Do NOT include tracked changes when sub-

mitting a MS on a CD or electronically.

4 AUTHOR(S)

When there are several authors, the covering letter

should indicate clearly which of them is responsible for

correspondence and, if possible, telephonically available

while the article is being processed. The contact address,

telephone number and email address should be men-

tioned if they differ from those given on the letterhead.

5 TITLE

The title should be as concise and as informative as

possible. In articles dealing with taxonomy or closely

related subjects, the family of the taxon under discus-

sion (see also 13.2) should be mentioned in brackets but

author citations should be omitted from plant names (see

also 13.6).

6 KEYWORDS

Up to 10 keywords (or index terms) should be pro-

vided in English in alphabetical sequence. The follow-

ing points should be borne in mind when selecting key-

words:

6.1 Keywords should be unambiguous, internatio-

nally acceptable words and not recently coined little-

known words.

6.2 They should be in a noun form and verbs should

be avoided.

6.3 They should not consist of an adjective alone;

adjectives should be combined with nouns.

6.4 They should not contain prepositions.

6.5 The singular form should be used for processes

and properties, e.g. evaporation.

6.6 The plural form should be used for physical

objects, e.g. augers.

6.7 Location (province and/or country); taxa (species,

genus, family) and vegetation type (community, veld

type, biome) should be used as keywords.

6.8 Keywords should be selected hierarchically

where possible, e.g. both family and species should be

included.

6.9 They should include terms used in the title.

6. 10 They should answer the following questions:

6.10.1 What is the active concept in the document (acti-

vity, operation or process).

6.10.2 What is the passive concept or object of the

active process (item on which the activity, operation or

process takes place).

6.10.3 What is the means of accomplishment or how is

the active concept achieved (technique, method, appara-

tus, operation or process).

6. 1 0.4 What is the environment in which the active con-

cept takes place (medium, location).

6.10.5 What are the independent (controlled) and depen-

dent variables?

Bothalia 42,2 (2012)

261

6.11 Questions 6.10.1 to 6.10.3 should preferably also

be answered in the title.

7 ABSTRACT

7. 1 An abstract of no more than 200 words should be

provided. Abstracts are of great importance and should

convey the essence of the article.

7.2 It should refer to the geographical area concerned

and, in taxonomic articles, mention the number of taxa

treated. It should not contain information not appearing

in the article.

7.3 In articles dealing with taxonomy or closely

related subjects all taxa from the rank of genus down-

wards should be accompanied by their author citations

(see also 13.6).

7.4 Names of new taxa and new combinations should

not be italicized but put in bold. If the article deals with

too many taxa, only the important ones should be men-

tioned.

8 TABLE OF CONTENTS

A table of contents should be given for all articles longer

than about 60 typed pages, unless they follow the strict

format of a taxonomic revision.

9 ACKNOWLEDGEMENTS

Acknowledgements should be kept to the minimum

compatible with the requirements of courtesy. Please

give all the initials of the person(s) you are thanking.

10 LITERATURE REFERENCES

In text

10.1 Literature references in the text should be cited as

follows: ‘Jones & Smith (1986) stated...’, or ‘...(Jones &

Smith 1986)’ or (Ellis 1988: 67) when giving a reference

simply as authority for a statement. For treatment of lit-

erature references in taxonomic papers see 14.

10.2 When more than two authors are involved in the

paper, use the name of the first author followed by et al.

10.3 When referring to more than one literature refer-

ence, they should be arranged chronologically and sepa-

rated by a semicolon, e.g. (Nixon 1940; Davis 1976;

Anon. 1981, 1984).

10.4 Titles of books and names of journals should

preferably not be mentioned in the text. If there is good

reason for doing so, they should be treated as described

in 10.12 and 10.13.

10.5 Personal communications are given only in the

text, not in the list of references. Please add the person’s

full initials to identify the person more positively, e.g. C.

Boucher pers. comm.

In References at end of article

10.6 References of the same author are arranged in

chronological sequence.

10.7 Where two or more references by the same author

are listed in succession, the author’s name is repeated

with every reference, except in an obituary, where the

name of the deceased in the list of publications (not in

the references) is replaced by an N-dash.

10.8 All publications referred to in the text, including

those mentioned in full in the treatment of correct names

in taxonomic papers, but no others, and no personal

communications, are listed at the end of the manuscript

under the heading References.

10.9 The references are arranged alphabetically

according to authors and chronologically under each

author, with a, b, c, etc. added to the year, if the author

has published more than one work in a year. This

sequence is retained when used in the text, irrespective

of the chronology.

10.10 If an author has published both on his own and

as a senior author with others, the solo publications are

listed first and after that, in strict alphabetical sequence,

those published with one or more other authors.

10.11 Author names are typed in CAPITAL LETTERS.

10.12 Titles of journals and of books are written out

in full and are italicized as follows: Transactions of the

Linnean Society of London 5: 171-217, or Biology and

ecology of weeds: 24.

10.13 Titles of books should be given as in Taxonomic

literature, edn 2 by Stafleu & Cowan and names of

journals as in the latest edition of World list of scientific

periodicals.

1 0. 14 Examples of references:

Collective book or Flora

BROWN, N.E. 1909. Asclepiadaceae. In W.T. Thiselton-Dyer, Flora

capensis 6,2: 518-1036. Reeve, London.

CUNNINGHAM, A.B. 1994. Combining skills: participatory

approaches in biodiversity conservation. In B.J. Huntley, Botanical

diversity in southern Africa. Strelitzia 1: 149-167. National Botanical

Institute, Pretoria.

Book

DU TOIT, A.L. 1966. Geology of South Africa, edn 3: 10-50. S.M.

Haughton (ed.). Oliver & Boyd, London.

HUTCHINSON, J. 1946. A botanist in southern Africa: 69. Gawthom,

London.

Journal

DAVIS, G. 1988. Description of a proteoid-restioid stand in Mesic

Mountain Fynbos of the southwestern Cape and some aspects of its

ecology. Bothalia 18: 279-287.

SMOOK, L. & GIBBS RUSSELL, G.E. 1985. Poaceae. Memoirs of

the Botanical Survey of South Africa No. 5 1 : 45—70.

STEBBINS, G.L. Jr. 1952. Aridity as a stimulus to plant evolution.

American Naturalist 86: 35M4.

262

Bothalia 42,2 (2012)

In press, in preparation

TAYLOR, H.C. in press. A reconnaissance of the vegetation of Rooib-

erg State Forest. Technical Bulletin, Department of Forestry.

VOGEL, J.C. 1982. The age of the the Kiiiseb river silt terrace at

Homeb. Palaeoecology of Africa 15. In press.

WEISSER, P.J., GARLAND, J.F. & DREWS, B.K. in prep. Dune

advancement 1937-1977 and preliminary vegetation succession chro-

nology at Mlalazi Nature Reserve, Natal, South Africa. Bothalia.

Thesis

KRUGER, F.J. 1974. The physiography and plant communities of the

Jakkalsrivier Catchment. M.Sc. (Forestry) thesis. University of Stel-

lenbosch.

MUNDAY, J. 1 980. The genus Monechma Hochst. (Acanthaceae tribe

Justiciae) in southern Africa. M.Sc. thesis. University of the Witwa-

tersrand, Johannesburg.

Miscellaneous paper, report, unpublished article, techni-

cal note, congress proceedings

ANON, no date. Eetbare plante van die Wolkberg. Botanical Research

Unit, Grahamstown. Unpublished.

BAWDEN, M.G. & CARROL, D.M. 1968. The land resources of

Lesotho. Land Resources Study No. 3, Land Resources Division,

Directorate of Overseas Surveys, Tolworth.

BOUCHER, C. 1981. Contributions of the Botanical Research Insti-

tute. In A.E.F. Heydom, Proceedings of workshop research in Cape

estuaries: 105-107. National Research Institute for Oceanology, CSIR,

Stellenbosch.

NATIONAL BUILDING RESEARCH INSTITUTE 1959. Report of

the committee on the protection of building timbers in South Africa

against termites, woodboring beetles and fungi, edn 2. CSIR Research

Report No. 169.

1 1 TABLES (also electronic submissions)

11.1 Each table should be presented on a separate

sheet and be assigned an Arabic numeral, i.e. the first

table mentioned in the text is marked ‘Table 1 ’.

11.2 In the captions of tables the word ‘TABLE’ is

written in capital letters. See recent numbers of Bothalia

for the format required.

1 1 .3 Avoid vertical lines, if at all possible. Tables can

often be reduced in width by interchanging primary hor-

izontal and vertical heads.

12 FIGURES (original or electronic submissions)

12.1 Line drawings (original artwork) should be in

jet-black Indian ink, on fine art paper, 200 gsm. Lines

should be clear enough to accommodate reduction. Do

not use draughtman’s film as it causes problems with the

scanning process.

12.2 Drawings in pencil will not be accepted.

12.3 Provide original drawings electronically as bit-

map TIF files, 600 dpi or preferably, 1200 dpi. Provide

photographs electronically as either TIF or JPG files,

600 dpi or higher. At the request of the Author, the Pub-

lications Section of SANBl will assist with the scan-

ning of original material. Figures should be planned to

fit, after reduction, into a width of either 80, 118 or 165

mm, with a maximum vertical length of 230 mm. Allow

space for the caption in the case of figures that will

occupy a whole page.

12.4 Graphs and histograms should be submitted

as stipulated in 3.9, or as TIF or JPG files at a resolu-

tion of 600 dpi or higher if generated in other pro-

grammes. Graphs and histograms generated in EXCEL

or MSWord, should be provided as is. File conversion

into the correct format will be aecommodated by SANBl

Graphics. Please do not supply embedded graphics in

the documents or files that are optimized for screen use.

Do not submit graphs and histograms in colour. If shad-

ing is used it should be easily discernible.

12.5 Photographs should be of excellent quality on

glossy paper with clear detail and moderate contrast so

that the figures can be seanned without retouching them

electronically. If submitted electronieally, provide as a

TIF or JPG file at 600 dpi or higher and not as a DOC,

PDF, EXCEL or POWERPOINT file.

12.6 Photograph mosaics should be submitted as sepa-

rate photographs or TIF/JPG files at 600 dpi or higher,

as well as a photocopy/layout of the mosaic. Final layout

of the mosaic will be done by our graphics department.

12.7 Do not number the original images but include a

scale bar. Indicate the lettering on the photocopy and not

on the original image or electronic copy.

12.8 If several illustrations are treated as components

of a single composite figure they should be designated

by eapital letters.

12.9 Note that the word ‘Figure’ should be written out

in full, both in the text and the captions and should begin

with a capital ‘F’ (but see 14.7 for taxonomic papers).

12.10 In the text the figure reference is then written as

in the following example: ‘The stamens (Figure 4A, B)

are...’

12.1 1 In captions, ‘FIGURE’ is written in capital letters.

12.12 Scale bars or scale lines should be used on fig-

ures.

12.13 In figures accompanying taxonomic papers,

voucher specimens should be given in the relevant cap-

tion.

12.14 Figures are numbered conseeutively with Ara-

bie numerals in the order they are referred to in the text.

These numbers, as well as the author’s name and an

indication of the top of the figure, must be written in soft

pencil on the back of all figures.

12.15 Captions of figures should not be pasted under

the photograph or drawing and should also not be

included in any electronic version of the figures.

12.16 Captions of figures should be collected together

and typed at the end of the MS and headed Captions for

figures.

Bothalia42,2 (2012)

263

12.17 Authors should indicate in pencil in the text

where they would like the figures to appear.

12.18 Authors wishing to have the originals of figures

returned must inform the editor in the original covering

letter and must mark each original ‘To be returned to

author’.

12.19 Authors wishing to use illustrations already pub-

lished elsewhere must obtain written permission before

submitting the manuscript and inform the editor of this

fact.

12.20 It is strongly recommended that taxonomic arti-

cles include dot maps as figures to show the distribution

of taxa. Maps will be reduced to column width (80 mm):

the symbols and numbers used must be large enough to

stand reduction. The maps should show: numbered grid

lines of latitude and longitude; the provinces of South

Africa; and a scale line. Maps of neighbouring countries

should be treated in the same way, with bordering states

clearly labelled. For orientation purposes, a small inset

map should appear in a comer of the figure.

12.21 ArcView GIS maps are aeceptable. The layout

representing all the appropriate themes (including grid

lines) should be submitted as an encapsulated postscript

file (EPS).

12.22 Colour figures are permitted only if: a) it will

clarify the article and b) the cost of reproduction and

printing is borne by the author.

12.23 Magnification of figures in the caption should be

given for the size as submitted.

13 TEXT

13.1 Asa rule, authors should use the plant names (but

not of all authors of plant names — see 13.6) as listed in

PRECIS (National Herbarium PREtoria Computerised

Information System).

13.2 Names of genera and infrageneric taxa are usu-

ally italicized, with the author citation (where relevant;

see 13.6) not italicized. Exceptions include names of

new taxa in the abstract, correct names given in the syn-

opsis or in paragraphs on species excluded from a given

supraspecific group in taxonomic articles; in checklists

and in indices, where the position is reversed, correct

names are not italicized and synonyms are italicized.

13.3 Names above generic level are not italicized.

13.4 In articles dealing with taxonomy, the complete

scientific name of a plant (with author citation) should

be given at the first mention in the text. The generic

name should be abbreviated to the initial thereafter,

except where intervening references to other genera with

the same initial could cause confusion (see 16.6).

13.5 In normal text, Eatin words are italicized, but in

the synopsis of a species, Eatin words such as nom. mid.

and et al. are not italicized (see 16.4, 17.2).

13.6 In accordance with Gamock-Jones & Webb

(1996) in Taxon 45: 285, 286, authors of plant names

are not to be added to plant names except in taxonomic

papers. Names of authors of plant names should agree

with the list published by the Royal Botanic Gardens,

Kew, entitled, Authors of plant names, edited by R.K.

Brummitt & C.E. Powell (1992).

13.7 Modem authors not included in the list should

use their full name and initials when publishing new

plant names. Other author names not in the list should be

in agreement with the recommendations of the Code.

13.8 Names of authors of publications are written

out in full, without initials, except in the synonymy in

taxonomic articles where they are treated like names of

authors of plant names.

13.9 Names of plant colleetors are italicized whenever

they are linked to the number of a specimen. The collec-

tion number is also italicized, e.g. Acocks 14407.

13.10 Surnames beginning with ‘De’, ‘Du’ or ‘Van’

begin with a capital letter unless preceded by an initial.

13.11 Eor measurements use only units of the Interna-

tional System of Units (SI). In taxonomic papers only

mm and m, should be used; in ecological papers cm or

m should be used.

13.12 The use of ‘±’ is preferred to c. or ca (see 17.8).

13.13 Numbers ‘one’ to ‘nine’ are spelt out in normal

text, and from 10 onwards they are written in Arabic

numerals.

13.14 In descriptions of plants, numerals are used

throughout. Write 2. 0^.5 (not 2^.5) and 2. 0-4. 5 6-9.

When counting members write 2 or 3 (not 2-3), but 2-A.

13.15 Abbreviations should be used sparingly but con-

sistently. No full stops are placed after abbreviations

ending with the last letter of the full word (e.g. edition =

edn; editor = ed.); after units of measure; after compass

directions; after herbarium designations; after countries,

e.g. USA and after well-known institutions, e.g. CSIR.

13.16 Apart from multi-access keys, indented keys

should be used with couplets numbered la- lb, 2a-2b,

etc. (without full stops thereafter).

13.17 Keys consisting of a single couplet have no num-

bering.

13.18 Manuscripts of keys should be presented as in the

following example:

la Leaves closely arranged on elongated stem; a submerged aquatic

with only capitula exserted ... lb. setaceum war. ptimilum

lb Leaves in basal rosettes; stems suppressed; small marsh plants,

ruderals or rarely aquatics:

2a Annuals, small, fast-growing pioneers, dying when habitat dries up;

capitula without coarse white setae; receptacles cylindrical:

3 a Anthers white .. .2. E. cinereum

3b Anthers black ... 3. E. nignm

2b Perennials, more robust plants; capitula sparsely to densely covered

with short setae:

13.19 Herbarium voucher specimens should be referred

to wherever possible, not only in taxonomic articles.

264

Bothalia 42,2 (2012)

13.20 The word Figure should be written out in full and

should begin with a capital F, also in captions where the

whole word is in capital letters (see 12.8-12.11).

1 4 SPECIES TREATMENT IN TAXONOMIC PAPERS

14.1 The procedure to be followed is illustrated in the

example (17.2), which should be referred to, because not

all steps are described in full detail.

14.2 The correct name (bold, not italicized) is to be

followed by its author citation (italicized) and the full

literature reference, with the name of the publication

written out in full (not italicized).

14.3 Thereafter all literature references, including

those of the synonyms, should only reflect author, page

and year of publication, e.g. C.E.Hubb. in Kew Bulletin

15: 307 (1960); Boris et al.: 14 (1966); Boris: 89 (1967);

Sims: t. 38 (1977); Sims: 67 (1980).

14.4 The description and the discussion should consist

of paragraphs commencing, where possible, with itali-

cized leader words such as flowering time, etymology,

diagnostic characters, distribution and habitat, with a

colon following the leader word and the first word of the

sentence beginning with a lower case letter.

14.5 When more than one species of a given genus is

dealt with in a paper, the correct name of each species

should be prefixed by a sequential number followed by

a full stop. Infraspecific taxa are marked with small let-

ters, e.g. lb., 12c., etc.

14.6 Names of authors are written as in 13.6, irrespec-

tive of whether the person in question is cited as the

author of a plant name or of a publication.

14.7 The word ‘figure’ is written as ‘fig.’, and ‘t.’ is

used for both ‘plate’ and ‘tablet’ (but see 12.9 for nonnal

text).

14.8 Literature references providing good illustrations

of the species in question may be cited in a paragraph

commencing with the word Illustrations followed by a

colon. This paragraph is given after the last paragraph of

the synonymy, see 17.2.

14.9 When new combinations are made, the full litera-

ture reference must be given for the basionym, e.g.:

Antimima saturata {L. Bolus) H.E.K. Hartmann,

comb. nov.

Ruschia saturata L. Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 122 (1929). Mesembryanthemum atrocinctum

N.E.Br.: 32 (1930). Type: Pillans BOL1R952 (BOL, holo.-photol).

14.10 Treatment of subspecies. Example:

37. Acacia robusta Burch., Travels in the interior

of southern Africa: 442 (1824); Harv.: 281 (1862); Oliv.:

349 (1871) pro parte excl. specim. IVelwitsch; Palmer

& Pitman: 807 (1973). Type: Cape Province, Kuruman

Dist. Takoon [Litakun], Burchell 2265 (K, holo.!).

(The species description encompassing the subspecific characteristics

and notes on distribution should follow here.]

Two subspecies are distinguished:

Leaf rachis glabrous or almost so; pods straight or slightly

curved 37a. subsp. robusta

Leaf rachis sparsely to densely pubescent; pods usually ± fal

cate 37b. subsp. clavigera

37a. subsp. robusta.

Brenan in Flora zambesiaca 3,1: 103 (1970); Ross: 37

(1971).

A. robusta Burch.: 442 (1824). [this is the basionym]

[The diagnostic description of the subspecies or reference to above key

and notes on distribution should follow here.]

37b. subsp. clavigera {E.Mey.) Brenan in Flora

zambesiaca 3,1: 104 (1970); Ross: 193 (1973). Type:

Natal, near Port Natal [Durban], Drege s.n. (K, iso.!; P,

Iragm.l).

A. clavigera E.Mey., Commentariorum 1: 168 (1836); Benth.: 510

(1875); Brenan: 365 (1958).

[The diagnostie deseription of the subspecies or reference to above key

and notes on distribution should follow here.]

15 CITATION OF SPECIMENS

15.1 Type specimen in synopsis: the following should

be given (if available): country (if not in RSA), prov-

ince, grid reference (at least for new taxa), locality as

given by original collector, modem equivalent of col-

lecting locality in square brackets (if relevant, e.g. Port

Natal [now Durban]), quarter-degree square, date of

collection (optional), collector’s name and collecting

number (both italicized).

15.2 The abbreviation s.n. (sine ntimero) is given after

the name of a collector who usually assigned numbers

to his collections but did not do so in the specimen in

question (see 15.11), or the herbarium number can then

be cited with no space between the herbarium and its

number e.g. Marloth SAM691 (see 17.2). The herbaria

in which the relevant type(s) are housed are indicated by

means of the abbreviations given in the latest edition of

Index Herbariorum.

15.3 The holotype (holo.) and its location are men-

tioned first, followed by a semicolon, the other herbaria

are arranged alphabetically, separated by commas.

15.4 Authors should indicate by means of an exclama-

tion mark (!) which of the types have been personally

examined.

15.5 If only a photograph, microfiche, electronic or

Aluka image was seen, write as follows: Anon. 422 (X,

holo.-BOL, photo.!), or Anon. 422 (X, holo.-Aluka

image, website accessed 14-08-2009), or (SDNH, holo.

e!).

15.6 Lectotypes or neotypes should be chosen for cor-

rect names without a holotype. It is not necessary to lec-

totypify synonyms.

15.7 When a lectotype or a neotype are newly chosen,

this should be indicated by using the phrase ‘here desig-

nated’ (see 17.2). If reference is made to a previously

Bothalia 42,2 (2012)

265

selected lectotype or neotype, the name of the designat-

ing author and the literature reference should be given. In

cases where no type was cited, and none has subsequently

been nominated, this may be stated as ‘not designated’.

15.8 In brief papers mentioning only a few species and

a few cited specimens the specimens should be arranged

according to the grid reference system: Provinces/countries

(typed in capitals) should be cited in the following order:

Namibia, Botswana, Limpopo (previously Northern Trans-

vaal, Northern Province), North-West (previously north-

eastern Cape and southwestern Transvaal), Gauteng (previ-

ously PWV), Mpumalanga (previously Eastern Transvaal),

Free State (previously Orange Free State), Swaziland,

KwaZulu-Natal (previously Natal), Lesotho, and Northern

Cape, Western Cape and Eastern Cape (Figure 1 ).

15.9 Grid references should be cited in numerical

sequence.

15.10 Locality records for specimens should preferably

be given to within a quarter-degree square. Records from

the same one-degree square are given in alphabetical

order, i.e (-AC) precedes (-AD), etc. Records from the

same quarter-degree square are arranged alphabetically

according to the collectors’ names; the quarter-degree

references must be repeated for each specimen cited.

15.11 The relevant international code of the herbaria in

which a collection was seen should be given in brackets

after the collection number; the codes are separated by

commas. The following example will explain the proce-

dure:

KWAZULU-NATAL.— 2731 (Louwsburg): 16 km E of Nongoma. (-

DD), 10-10-1960, Reiser 354 (BM, K, PRE); near Dwarsrand, Van der

Merwe 4789 (BOL, M). 2829 (Harrismith): near Groothoek, (-AB),

Smith 234\ Koffiefontein, (-AB), Taylor 720 (PRE); Cathedral Peak

Forest Station, (-CC), 8 July 1905, Marriot s.n. (KMG); Wilgerfon-

tein, Roia 426. Grid ref unknown: Sterkstroom, Sttydom 12 (NBG).

15.12 For records from outside southern Africa authors

should use degree squares without names, e.g.:

KENYA. — 0136: Nairobi plains beyond race course, Napier 485.

15.13 Monographs and revisions: in the case of all major

works of this nature it is assumed that the author has

investigated the relevant material in all major herbaria and

that he has provided the specimens seen with determinavit

labels. It is assumed further that the author has submitted

distribution maps for all relevant taxa and that the distribu-

tion has been described briefly in words in the text. Under

the heading ‘Vouchers’ no more than five specimens

should be cited, indicating merely the collector and the

collector’s number (both italicized). Specimens are alpha-

betically arranged according to collector’s name. If more

than one specimen by the same collector is cited, they are

arranged numerically and separated by a comma. A collec-

tor’s name and the voucher number(s) is separated from

the next collector by a semicolon. The purpose of the cited

specimens is not to indicate distribution but to convey the

author’s concept of the taxon in question.

15.14 The herbaria in which the specimens are housed

are indicated by means of the abbreviation given in the

latest edition of Index Herbariorum. They are given

between brackets, arranged alphabetically and separated

by commas behind every specimen as in the following

example:

Vouchers: Arnold 64 (PRE); Fisher 840 (NH, NU, PRE); Flanagan

831 (GRA, PRE), 840 (NH, PRE); Marloth 4926 (PRE, STE); Schelpe

6161, 6163, 6405 (BOL); Schlechter 4451 (BM, BOL, GRA, K, PRE).

15.15 If long lists of specimens are given, list them

together before Acknowledgements under the head-

ing Specimens examined. They are arranged alphabeti-

cally by the collector’s name and then numerically for

each taxon. The species is indicated in brackets by the

number that was assigned to it in the text and any infra-

specific taxa by a small letter; this number follows the

specimen number. If more than one genus is dealt with

in a given article, the first species of the first genus men-

tioned is indicated as 1.1. This is followed by the inter-

national herbarium designation. Note that the name of

the collector and the collection number are italicized:

Acocks 14724 (1.13a) BOL, K, P; 12497 (2.1b) BM, K, PRE. Archer

7507(1.4) BM, G.

Barker 9738 (1) NBG; 1916 (2) NBG; 295, 4766, 9478, 9796, 10330

(4) NBG; 1919 (5) BOL, NBG; 1917,1923, 1935, 2570, 2606, 2646,

3332, 4198, 4858, 10534, 10801 (5) NBG. Burchell 2847 (2.8c) MB,

K. Biirman 2401 (3.3) MO, S. B.L. Burn 789 (2.6) B, KMG, STE.

Esterhuysen 77497(1) BOL; 1433 (5) BOL; 71402 (5) NBG.

16 SYNONYMS

16.1 In a monograph or a revision covering all of

southern Africa, all synonyms based on types of south-

ern African origin, or used in southern African literature,

should be included.

16.2 Illegitimate names are designated by nom. illeg.

after the reference, followed by non with the author and

date, if there is an earlier homonym.

16.3 Nomina nuda {nom. nnd.) and invalidly published

names are excluded unless there is a special reason to

cite them, for example if they have been used in promi-

nent publications.

16.4 In normal text, Latin words are italicized, but in

the synopsis of a species Latin words such as nom. mid.,

et al. are not italicized (see 13.5, 17.2).

16.5 Synonyms should be arranged chronologically

into groups of nomenclatural synonyms, i.e. syno-

nyms based on the same type, and the groups should be

arranged chronologically by basionyms, except for the

basionym of the correct name which is dealt with in the

paragraph directly after that of the correct name.

16.6 When a generic name is repeated in a given syn-

onymy it should be abbreviated to the initial, except

where intervening references to other genera with the

same initial could cause confusion (see 13.4).

17 DESCRIPTION AND EXAMPLE OF SPECIES

TREATMENT

17.1 Descriptions of all taxa of higher plants should,

where possible, follow the sequence: Habit; sexuality;

underground parts (if relevant). Indumentum (if it can be

266

Bothalia42,2 (2012)

easily described for the whole plant). Stems/branches.

Bark. Stipules. Leaves: arrangement, petiole absent/

present, pubescence; blade: shape, measurements, apex,

base, texture, colour; margin; midrib: above/below;

petiole. Inflorescence: type, shape, measurements, posi-

tion; bracts/bracteoles, involucral bracts: inner, outer.

Flowers: shape, measurements, sex, colour. Receptacle.

Calyx. Corolla. Disc. Androecium. Gynoecium. Fruit.

Seeds (apply the same sequence of relevant features as

in flowers). Flowering time. Chromosome number (ref-

erence). Conservation status. Figure number (word writ-

ten out in full).

1 7.2 Example (not factual for this species):

1. Englerophytum magalismontanum (Sond.)

T.D.Penn., The genera of Sapotaceae: 252 (1991). Type:

Gauteng, Magaliesberg, Zeyher 1849 (S, holo.-BOL,

photo.!).

Bequaertiodendron magalismontanum (Sond.) Heine & Hemsl.: 307

(1960); Codd: 72 (1964); Elsdon: 75 (1980).

ChysophyUum magalismontanum Sond.: 721 (1850); Harv.: 812

(1867); Engl.: 434 (1904); Bottmar: 34 (1919). Zeyherella magalis-

montana (Sond.) Aubrev. & Pellegr.: 105 (1958); Justin: 97 (1973).

ChysophyUum argyrophyllum Hiem: 721 (1850); Engl.: 43 (1904).

Boivinella argyrophylla (Hiem) Aubrev. & Pellegr.: 37 (1958); Justin

et al.: 98 (1973). Types: Angola, Welwitsch 4828 (BM, lecto.!, here

designated; PRE!); Angola, Welwitsch s.n. (BM!).

Chtysophyllum wilmsii Engl.: 4, t. 16 (1904); Masonet: 77 (1923);

Woodson: 244 (1937). Boivinella wilmsii (Engl.) Aubrev. & Pellegr.:

39 (1958); Justin: 99 (1973). Type: without locality and collector [B,

holo.t; Kl, P, lecto. e!, designated by Aubrev. & Pellegr.: 38 (1958),

PRE!,S!,W!,Z!].

Bequaertiodendron fruticosa De Wild,: 37 (1923), non Bonpl.: 590

(1823); D.Bakker: 167 (1929); H.Fr.: 302 (1938); Davy: 640 (1954);

Breytenbach: 117 (1959); Clausen: 720 (1968); Palmer: 34 (1969),

Type: Mpumalanga, Tzaneen Dist., Granville in Herb. Pillans K48625

(K, holo.!; G!, P!, PREI, S!-Aluka image, website accessed 14-08-

2009).

B. fragrans auct. non Oldemann: Glover: 149, t. 19 (1915); Henkel:

226 (1934); Stapelton: 6 ( 1 954).

Illustrations: Harv.: 812 (1867); Henkel: t. 84 (1934?); Codd: 73

(1964); Palmer: 35 (1969).

Woody perennial; main branches up to 0.4 m long, erect

or decumbent, grey woolly-felted, leafy Bark smoothish,

pale, beige-brown, flaking; lenticellate, splitting longi-

tudinally. Stipules paired, spines up to 50 x 45 x ± 2 mm,

unmistakeably large and flattened, roughly triangular in

outline, the flared basiscopic lobe margins folded over in

larger stipules, spines held at ± 90° to stem. Stem erect,

simple or 1- or 2-branched, either from base or from upper

stem nodes. Leaves linear to oblanceolate, 3-10(-23)

X 1.0-1.5(-4.0) mm, obtuse, base broad, half-clasping.

Lleads/Inftorescence heterogamous, campanulate, 7-8 x

5 mm, solitary, sessile at tip of axillary shoots; involucral

bracts in 5 or 6 series, inner exceeding flowers, tips subo-

paque, white, very acute. Receptacle nearly smooth. Flow-

ers ± 23-30, 7-11 male, 16-21 bisexual, yellow, tipped

pink. Stamens unilateral and declinate; filaments 7-9 mm

long, exserted for 7-8 mm; anthers 3^ mm long, dull pink.

Ovary ovoid, ± 3.5 mm long; style dividing near apex of

anthers, style branches 3-4 mm long, recurved. Pappus

bristles very many, equalling corolla, scabridulous. Cap-

sules/achenes subglobose, 3-lobed, 6-7 mm long. Seeds

tetrahedral, colliculate, ± 2 mm long. Flowering time:

Septemher. Chromosome number: 2n = 22. Figure 23B.

17.3 As a rule, shape should be given before measure-

ments.

17.4 In general, if an organ has more than one of the

parts being described, use the plural, otherwise use the sin-

gular, for example, petals of a flower but blade of a leaf

17.5 Language must be as concise as possible, using

participles instead of verbs.

17.6 Dimension ranges should be cited as in 17.2.

17.7 Care must be exercised in the use of dashes and

hyphens. A hyphen is a short stroke joining two sylla-

bles of a word, e.g. ovate-lanceolate or sea-green, with

no space between the letter and the stroke. An N-dash

(en) is a longer stroke commonly used instead of the

word ‘to’ between numerals, ‘2-5 mm long’ (do not use

it between words but rather use the word ‘to’, e.g. ‘ovate

to lanceolate’; it is produced by typing three hyphens

with spaces in between, or in MS Word the code is alt +

0150. An M-dash (em) is a stroke longer than an N-dash

and is used variously, e.g. in front of a subspecific epi-

thet instead of the full species name; it is produced

by typing two hyphens with spaces in between, or in

MSWord the code is alt + 0151. See also 2.10-2.12.

1 7.8 The use of ‘±’ is preferred to c. or ca when describ-

ing shape, measurements and dimensions (see 13.12).

17.9 The decimal point replaces the comma in all units

of measurement, e.g. leaves 1.0-1. 5 mm long.

18 NEWTAXA

18.1 The name of a new taxon must be accompanied

by at least a Latin diagnosis. Authors should not pro-

vide full-length Latin descriptions unless they have the

required expertise in Latin at their disposal.

18.2 It is recommended that descriptions of new taxa

be accompanied by a good illustration, preferably a line

drawing, or a photograph (second choice) and a distri-

bution map.

18.3 Example:

109. Helichrysum jubilatum Hilliard, sp. nov., H. alsi-

noidei DC. affinis, sed foliis ellipticis (nec spatulatis),

inflorescentiis compositis a foliis non circumcinctis, flo-

ribus femineis numero quasi dimidium hermaphrodito-

rum aequantibus (nec capitulis homogamis vel floribus

femineis 1-3 tantum) distinguitur.

Herba annua e basi ramosa; caules erecti vel decum-

bentes, 100-250 mm longi, tenuiter albo-lanati, remote

foliati. Folia plerumque 8-30 x 5-15 mm, sub capitu-

lis minora, elliptica vel oblanceolata, obtusa vel acuta,

mucronata, basi semi-amplexicauli, utrinque cano-

lanato- arachnoidea. Capitula heterogama, campanu-

lata, 3.5- 4.0 x 2.5 mm, pro parte maxima in paniculas

cymosas tenuinales aggregata; capitula subterminalia

interdum solitaria vel 2 vel 3 ad apices ramulorum nudo-

rum ad 30 mm longorum. Bracteae involucrales 5-seri-

atae, gradatae, exteriores pellucidae, pallide stramineae,

dorso lanatae, seriebus duabus interioribus subaequali-

bus et flores quasi aequantibus, apicibus obtusis opacis

Bothalia42,2 (2012)

267

niveis vix radiantibus. Receptaculum fere laeve. Flores

± 35-41. Achenia 0.15 mm longa, pilis myxogenis prae-

dita. Pappi setae multae, corollam aequantes, apicibus

scabridis, basibus non cohaerentibus.

TYPE. — Northern Cape, 2817 (Vioolsdrif): Richters-

veld, (-CC), ± 5 miles E of Lekkersing on road to Stink-

fontein, kloof in hill south of road, annual, disc whitish,

7-11-1962, Nordenstam 1823 (S, holo.; E, NH, PRE).

19 PROVINCES OF SOUTH AFRICA (Oct 1996)

FIGURE 1. — 1, Western Cape; 2, Eastern Cape; 3, Northern Cape; 4,

Free State (previously Orange Free State); 5. KwaZulu-Natal

(previously Natal); 6, North-West (previously northeastern Cape

and southwestern Transvaal); 7, Gauteng (previously PWV); 8,

Mpumalanga (previously Eastern Transvaal); 9, Limpopo (previ-

ously Northern Transvaal. Northern Province).

20 PROOFS

Only page proofs are normally sent to authors. They

should be corrected in red ink and be returned to the edi-

tor as soon as possible. Do not add any new information.

21 REPRINTS

Reprints will no longer be issued. A PDF file of the

article will be sent via the E-mail to authors and co-

authors. It is for private use only, the SANBI copyright

protects it from being used in another publication.

22 DOCUMENTS CONSULTED

Guides to authors of the following publications were

made use of in the compilation of the present guide:

Annals of the Missouri Botanic Garden, Botanical Jour-

nal of the Linnean Society, Flora of Australia, Smithso-

nian Contributions to Botany, South African Journal of

Botany (including instructions to authors of taxonomic

papers). South African Journal of Science.

23 ADDRESS OF PRODUCTION EDITOR

Manuscripts should be submitted to: Yolande

Steenkamp, Production Editor: Bothalia, South Afri-

can National Biodiversity Institute, Private Bag XlOl,

Silverton 0184; or, preferably, e-mailed to:

y.steenkamp(^sanbi.org.za.

24 FSA CONTRIBUTIONS

24.1 Figures and text must conform to Bothalia for-

mat.

24.2 These articles will be considered as a full contri-

bution to the Flora of southern Africa and will be listed

as published in the "Plan of Flora of southern Africa',

which appears in all issues of the FSA series.

25 PLACE NAMES

Ensure that local place names are correct. If in doubt,

consult the Internet at

http://sagns.dac.gov.za/searchplacenamedatabase.asp

INDEX

abbreviation, 13.4, 13.15, 15.2, 15.14, 16.6

abstract, 2.1, 2.3, 2.4, 7, 13.2

acknowledgements, 2.4, 9

address of

authors, 2.3, 4

editor, 23

alphabetical, 6, 10.9, 10.10, 15.3, 15.10, 15.13, 15.14, 15.15

Aluka image, 15.5, 17.2

Arc View GIS maps, 12.21

Arabic numerals, 11.1, 12.14, 13.13

author(s), 1.3, 2.1, 2.3, 4. 10.14, 12.17-12.19

address, 2.3, 4

citation, 5, 7.3, 13.2, 13.4, 14.2

first, 10.2

names, 2.3. 10.2, 10.7, 10.9, 10.11, 12.14, 13.7, 13.8, 14.3, 14.6, 15.7

names of plant names, 5, 13.1, 13.2, 13.6, 13.7, 13.8, 14.6

senior, 10.10

basionym, 14.10

book reviews, 1.1, 2.4

books, 10.4, 10.12, 10.13, 10.14

Bothalia, 1,2.2, 11.2,24.1

brief taxonomic articles. 15.8

BRUMMITT, R.K. & POWELL, C.E. (eds) 1992. Authors of plant

names. Royal Botanic Gardens, Kew, 13.6

c., 13.12, 17.8

ca, 13.12, 17.8

Cape, 15.8, 18.3, 19

capital letters, 10.11, 11.2, 12.8, 12.9, 12.11, 13.20, 15.8

captions, 2.1, 2.4, 2.5, 11.2, 12.3, 12.9, 12,11-12.13, 12.15, 12.16,

12.23, 13.20

CD, 3.2, 3.8, 3.11

CDR file, 3.9

checklist, 13.2

chromosome number, 17.1, 17.2

chronological sequence, 10.3, 10.6, 10.9, 16.5

citation

author, 5, 7.3, 13,2. 13.4, 14.2

of specimens, 15

cm, 13.1 1

collection

date, 15.1

number, 13.9, 15.1, 15.2, 15.11, 15.13, 15.15

collective book, 10.14

collector, 13.9, 15.1, 15.2, 15.10, 15.13, 15.15

colon, 2.13, 14.4, 14.8

colour figures, 12.22

comma, 2.13, 15.3, 15.11, 15.13, 15.14, 17.9

compass directions, 13.15

composite figure, 12.8

congress proceedings, 10.14

contents, 8

CorelDraw up to version 14, 3.9

correspondence, 4

countries, 6.7, 15.8

deadline dates for submission of MS, 1.6

268

Bothalia42,2 (2012)

decimal point, 17.9

description and example of species treatment, 17

determinavit labels, 15.13

dimension ranges, 1 7,6

discussion, 2.4, 14.4

distribution maps, 12.20, 15.13, 18.2

DOC, 12.5

documents consulted, 22

dot maps, 12.20-12.23, 15.13, 18.2

double

line spacing, 2.1

space, 2.13

dpi (dots per inch), 12.3-12.6

drawing paper, 12.1

drawings, 3.7, 12.1-12.3

Eastern Transvaal, see Mpumalanga, 15.8, 19

e!, 15.5, 17.2

edition, 13.15

editor, 1.5, 12.18, 13.15,23

editorial

board, 1.4

policy, 1

electronic

copy, 12.7

files, 3, 3.2, 3.8-3.10, 12.3-12.6

image, 15.5, 17.2

submissions of graphics, tables, 11,12

E-mail, 3.2, 3.8, 21

encapsulated postscript file (EPS), 3.9, 12.21

EPS file, 3.9, 12.21

etal., 10.2, 13.5, 14.3, 17.2

example of

new taxa, 18.3

species treatment, 17.2

exclamation mark, 2.13, 15.4

EXCEL file, 12.4, 12.5

family name, 5, 6.7

fig., 14.7

figure(s), 12, 13.20, 14.7, 17.1

colour, 12.22

electronic submissions of, 12

reduction of, 12.1, 12.3, 12.20

returned, 12.18

file

DOC, 12.5

electronic, 3, 3.2

extensions, 3.9

CDR, 3.9

EPS, 3.9, 12.21

EXCEL, 12.4, 12.5

JPG,

PDF, 12.5,21

POWERPOINT, 12.5

RTF, 3.3

TIF, 3.9, 12.3-12.6

firewall, 3.8

first author, 10.2

first language, 1 .3

flora, 10.14

Flora of southern Africa, 24

footnote, 3.4

Free State (previously Orange Free State), 15.8, 19

FSA contributions, 24

FTP site, 3.8

full stop, 2.13, 13.15, 13.16, 14.5

GARNOCK-JONES, P.J. & WEBB, C.J. 1996. The requirement to cite

authors of plant names in botanical journals. Taxon 45: 285, 286,

13.6

Gauteng (previously PWV), 15.8, 17.2, 19

genera, 13.2

generic name, 13.3, 13.4, 16.6

geographical area, 7,2

granting agencies, 2.3

graphics, 3,7

electronic submissions of, 12

graphs, 3.7, 1 2.4

grid reference system, 15.1, 15.8, 15.9, 15.11

headings, 3.4 3.6

sequenee of, 2.3, 2.4

herbaria, 15.2, 15.3, 15.11, 15.13, 15,14

herbarium

code, 15.11

designations, 13.15, 15.15

numbers, 15.2

voucher specimens, 12.13, 13.19

here designated, 15.7, 17.2

histograms, 12.4

holo., 14.10, 15.3, 15.5, 17.2, 18.3

holotype, 15.3, 15.6

homonym, 16.2

hyphenated words, 2.9

hyphen, 2.10-2.12, 17.7

illegitimate names (nom. illeg.), 16.2

illustrations, 12.1-12.3, 12.8, 12.19, 14.8, 17.2

previously published, 12.19

image files, 3.8-3. 1 0

indentations, 3.4, 3.6

Index Herbariorum, 15.2, 15.14

index of names, 2.4

indices, 13.2

infrageneric taxa, 13.2

initials, 9, 10.5, 13.7

in prep., 10.14

in preparation, 10.14

in press, 10.14

International

Code of Botanical Nomenclature, 13.7

System of Units (SI), 13.11

invalidly published names, 16.3

italics, 3.5, 7.4, 10.12, 13.2, 13.3, 13.5, 13.9, 14.2, 15.1, 15.13, 15.15

journals, 10.4, 10.12, 10.14

names of, 10,4, 10.13

JPG file, 12.3-12.6

justify, 2.8

keys, 2.16, 3.4, 13.16, 13.17, 13.18, 14.10

keywords, 2,3, 2.4, 6

KwaZulu-Natal (previously Natal), 15.8, 19

language, 1 .3

Latin, 13.5, 16.4

descriptions, 18.1

layout, 2.2, 12.6

lecto., 15.6, 15.7, 17.2

lectotype, 15.6, 15.7, 17.2

Limpopo (previously Northern Transvaal, Northern Province), 15.8, 19

line

drawings, 2.1, 12.1, 18.2

spacing, 2.1

literature

references, 2.1, 10, 14.2, 14.3, 14.8, 14.9

within synonymy, 14.8

localities outside southern Africa, 15.12

locality, 15.1, 15.10

location, 6.7

m, 13.11

magnification of figures, 12.12, 12.23

manuscript

language, 1.3, 17.5

requirements, 1.1, 1.2, 1.3,2

.sequence, 2.3, 2.4

map

Arc View GIS, 12.21

distribution, 12.20, 15.13, 18.2, 19

dot, 12.20

neighbouring countries, 12.20

M-dash, 2.12, 17.7

mm, 13.11

margin, 2.1, 2.16, 3.4, 17.1

material, 2.3, 2.4

measurements, 13.11, 17.1, 17.3, 17.8, 17.9

methods, 2.4, 6. 10.3

microfiche, 15.5

miscellaneous paper, 10.14

monograph, 2.4, 15.13, 16.1

Mpumalanga (previously Eastern Transvaal), 15.8, 19

MSWord, 2.7, 2. 1 1 , 2. 1 2, 2. 1 5, 3.3, 3.5, 1 7.7

name(s)

collector’s, 15.10

illegitimate, 16.2

Bothalia 42,2 (2012)

269

invalidly published, 16.3

of author(s), 2.3, 10.2, 10.7, 10.9, 10.11, 12.14, 13.7, 13.8, 14.3,

14.6, 15.7

name(s) (cont.)

of authors of plant names, 5, 7.3, 13.1, 13.2, 13.6, 13.7, 13.8, 14.6

of publications, 13.8

Natal, see KwaZulu-Natal, 15.8, 19

N-dash,2.11, 10.7, 17.7

neighbouring countries, 12.20

neotype, 15.6, 15.7

new

combinations, 7.4, 14.9

taxa, 7.4, 13.2, 13.7, 15.7, 18

nom. illeg., 16.2

nom. nud., 13.5, 16.3, 16.4

non-breaking space, 2.7

normal style, 3.1

Northern

Province, see Northern Transvaal, 15.8, 19

Transvaal, see Northern Province, Limpopo, 15.8, 19

North-West. 15.8, 19

notes, 1, 2.4

technical, 10.14

number

chromosome, 17.1, 17.2

herbarium, 15.2

numbering, 12.7, 13.13

of figures, 12.7, 12.14, 17.1

ofkeys, 13.16, 13.17

of pages, 2.5

of taxa, 14.5, 15.15

numerals, Arabic, 11.1, 12.14

obituaries, 1.1, 2.4, 10.7

Orange Free State, see Free State, 15.8, 19

original drawings/figures, 12.1-12.3, 12.7, 12.18

page charges, 1.5

paragraghs, 3.6

PDF file, 12.5,21

pencil drawings, 12.2

permission (written)

to copy material, 3.8

to use previously published material (text and illustrations), 12.19

pers. comm., 10.5, 10.8

personal communications (pers. comm.), 10.5, 10.8

photocopies, 2.1, 12.6, 12.7

photograph, 3.7, 12.3, 12.5, 12.6, 12.15, 15.5, 18.2

mosaic, 12.6, 12.7

plant

collectors, 13.9

name, 5, 13.4, 13.6, 13.7, 13.8, 14.6

plate (t.), 14.7

POWERPOINT fde, 12.5

PRECIS (National Herbarium PREtoria Computerised Information

System), 13.1

prepositions, 6.4

proceedings, 10.14

proofs, 20

provinces, 6.7, 15.1, 15.8

of South Africa, 15.8, 19

publications, 10.8, 13.8, 14.3

name of, 14.2

solo, 10.10

year of, 10.9, 14.3

PWV, see Gauteng, 15.8, 19

quarter-degree squares, 15.1, 15.10

quotes, 2.15

reduction of

figures, 12.1, 12.3, 12.20

tables, 11.3

referees, 1.4

reference(s), 2.4, 2.5, 10.6, 10.7-10.9, 10.14

figure, 12.10

grid, 15.1, 15.8, 15.9, 15.11

list, 10.5, 10.8, 10.9

literature, 2.1, 10

report. 10.14

reprints, 2 1

requirements for manuscript, 2

results, 2.4

revision, 2.4, 8, 15.13, 16.1

RTF file, 3.3

scale bar, 12.7, 12.12

scan/scanning, 12.1, 12.3, 12.5

semicolon, 2.13, 10.3, 15.3, 15.13

senior author, 10.10

sequence of headings, 2.3, 2.4

short notes, 1.1, 2.4

space

double, 2.13

non-breaking, 2.7

one, 2.16

special characters, 2.6

species treatment in taxonomic papers, 14.1-14.9

specimens examined, 2.4, 15.15

square brackets, 15.1, 17.2

STAFLEU, F.A. & COWAN, R.S. 1976-1988. Taxonomic literature.

Vols 1-7, 10.13

style(s), 3.1, 3.4

submission of MS, 1.2, 1.6

subspecies treatment in taxonomic papers, 14.10

surnames, 13.10

synopsis, 13.2, 13.5, 15.1, 16.4

synonymy, 13.8, 14.8, 16.6

t., 14.3, 14.7, 17.2

table(s), 2.1, 2.4, 2.5, 11

electronic submissions, 1 1

of contents, 8

tablet (t.), 14.7

tabs, 3.4

taxa

name of, 5, 7.4, 10.8, 13.2, 13.3

new, 7.4, 13.2, 13.7, 15.7, 18

numbering of, 14.5, 15.15

taxonomic

articles/papers, 7.2, 10.8, 12.13, 12.20, 13.2, 13.6, 13.8, 13.11, 14

revision, 8

treatment

species, 14.1-14.9

subspecies, 14.10

taxonomy, 5, 7.3, 13.4

technical note, 10.14

text, 2.1, 3.7, 10.1, 10.4, 10.5, 10.8, 10.9, 11.1, 12.9, 12.10, 12.14,

12.17, 13, 15.13, 15.15, 16.4

thesis, 10.14

TIF file, 3.9, 12.3-12.6

times sign, 2.14

title, 2.3, 5, 6.9, 6.11

ofbooks, 10.4, 10.12, 10.13, 10.14

ofjoumals, 10.4, 10.12, 10.13, 10.14

page, 2.3, 2.5

tracked changes, 3.11

Transvaal, 15.8, 17.2, 19

type, 15.2, 15.4, 15.7, 16.1, 16.5, 17.2

here designated, 15.7, 17.2

not designated, 15.7

specimen, 15.1

units of measure, 13.11, 13.15, 17.9

unpublished article, 10.14

voucher(s) specimens, 12.13, 13.19, 15.13, 15.4

World list of scientific periodicals, 10.13

year of publication, 10.9, 14.3

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CONTENTS

1. The emerging invasive alien plants of the Drakensberg Alpine Centre, southern Africa. C.

CARBUTT 71

2. Systematics of the southern African genus Ixia (Iridaceae: Crocoideae): 4. Revision of sect.

D/c/7o/7e. P. GOLDBLATT and J.C. MANNING 87

3. Systematics of the hypervariable Moraea tripetala complex (Iridaceae: Iridoideae) of the southern

African winter rainfall zone. P. GOLDBLATT and J.C. MANNING Ill

4. New species and subspecies of Babiana, Hesperantha, and Ixia (Iridaceae; Crocoideae) from

southern Africa; range extensions and morphological and nomenclatural notes on Babiana

and Geissorhiza. P. GOLDBLATT and J.C. MANNING 137

5. A taxonomic revision of the southern African native and naturalized species of Silene L.

(Caryophyllaceae). J.C. MANNING and P. GOLDBLATT 147

6. Notes on African plants:

Cyperaceae. Identity and typification of Carex cognate and status of C. drakensbergensis. C.

ARCHER and K. BALKWILL 190

Fabaceae. Ophrestia oblongifolia (Phaseoleae) in southern Africa. A.N. MOTEETEE and B.-

E. VANWYK 187

Hyacinthaceae. Albuca teniiifolia and A. shawii (Omithogaloideae), two distinct species from

South Africa. M. MARTINEZ- AZORIN, M.B. CRESPO and A.P. DOLD 193

Iridaceae. Notes on Moraea subg. Visciramosa (Iridoideae), including the new species Moraea

saldanhensis from the Atlantic coast of South Africa, recognition of M. inconspicua

subsp. namaqnensis, and pollination biology in M. riviilicola. P. GOLDBLATT and

J.C. MANNING 210

Iteaceae. Taxonomic and nomenclatural notes on the genus Choristylis. M. JORDAAN .... 196

Nyctaginaceae. Taxonomic status of Commicarpus fruticosus. M. STRUWIG and S.J.

SIEBERT 201

Poaceae. The taxonomic and conservation status of Agrostis eriantha var. planifolia. J.E.

VICTOR, A.C. MASHAU and V.J. NGOBENI 202

Rubiaceae. Taxonomic notes on Sericanthe andongensis and a new combination and status in

Sericanthe from Limpopo, South Africa. M. JORDAAN and H.M. STEYN 204

7. Nomenclature and typification of southern African species of Euphorbia. P.V BRUYNS 217

8. ESA contributions 20: Asteraceae: Anthemideae: Inezia. N. SWELANKOMO 247

9. South African National Biodiversity Institute: publications 1 January 2011 to 31 December 201 1.

Compiler; Y. STEENKAMP 251

10. Guide for authors to Bothalia 259

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