Volume 44 = Number 1

Provided by SANBI

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Michelle Hamer

Director: Zoological Systematics,

South African National

Biodiversity Institute, South Africa

Associate Editors and

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Stephen M. Awoyemi

Religion and Conservation

Research Collaborative, Nigeria

Saliem Fakir

WWE South Africa, South Africa

Mathieu Rouget

University of KwaZulu-Natal,

South Africa

Dave Balfour

Independent consultant,

Eastern Cape, South Africa

John Manning

South African National

Biodiversity Institute, South Africa

Krystal Tolley

South African National

Biodiversity Institute, South Africa

rast Page i of il Index a a aaa a a a ee SAE ae eo ae

Bothalia

AFRICAN BIODIVERSITY & CONSERVATION

ISSN: 0006-8241 (print)

ISSN: 2311-9284 (online)

Information for Authors and Readers

Original Research

A new long-tubed subspecies of Pelargonium dipetalum (section Hoarea) (Geraniaceae) from the Albertinia-

Swellendam area in Western Cape Province, South Africa

M. Marianne le Roux, John C. Manning

: Review Article

Factors influencing the adaptation and distribution of Colophospermum mopane in southern Africa’s mopane

savannas — A review

Rudzani A. Makhado, Isaac Mapaure, Martin J. Potgieter, Wilmien J. Luus-Powell, Amant T. Saidi

| Short Note

Moraea orthrosantha (Iridaceae: Irideae), a new species from Namaqualand, South Africa

Peter Goldblatt, John C. Manning

A new species of Berkheya (Asteraceae, Arctotideae) from the Northern Cape, South Africa

Nicola G. Bergh, Nick A. Helme

Taxonomic status of Pelargonium reniforme Curt.

Janine E. Victor, Mmamphe Aphane

Ledebouria caesiomontana A.J. Hankey & N.Hahn (Hyacinthaceae: Hyacinthoideae): A new species from the

Blouberg centre of endemism, Limpopo, South Africa

Andrew J. Hankey, Norbert Hahn, Matt H. Buys

Gladiolus filiformis, a poorly known species from North West Province, South Africa

Norbert Hahn, Hermuen Roux

_ Lectotypification of Kniphofia pauciflora Baker (Asphodelaceae: Asphodeloideae)

Himansu Baijnath, Syd Ramdhani

Nomenclatural adjustments in African plants 1

Peter Goldblatt, John C. Manning

Schoenefeldia transiens (Poaceae): Rare new record from the Limpopo Province, South Africa

Aluoneswi C. Mashau, Albie R. Gétze

Reviewer Acknowledgement

Reviewers who contributed to Volume 44, Number 1

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nbKOtENASKaLONM Oke NUVI GUANEL ANSLEY CESS)

- Page 1 of 8

A new long-tubed subspecies of Pelargonium

dipetalum (section Hoarea) (Geraniaceae) from

the Albertinia-Swellendam area in Western Cape

Authors:

M. Marianne le Roux!?

John C. Manning**

Affiliations:

1National Herbarium, South

African National Biodiversity

Institute, South Africa

*Department of Botany and

Plant Biotechnology, University

of Johannesburg, South Africa

3Compton Herbarium, South

African National Biodiversity

Institute, South Africa

4Research Centre for Plant

Growth and Development,

University of KwaZulu-Natal,

South Africa

Correspondence to:

Marianne le Roux

Email:

m.leroux@sanbi.org.za

Postal address:

Private Bag X101, Silverton

0184, South Africa

Dates:

Received: 19 May 2014

Accepted: 17 Sept. 2014

Published: 08 Dec. 2014

How to cite this article:

Le Roux, M.M. & Manning,

J.C., 2014, ‘A new long-tubed

subspecies of Pelargonium

dipetalum (section Hoarea)

(Geraniaceae) from the

Albertinia-Swellendam area in

Western Cape Province, South

Africa’, Bothalia 44(1), Art.

#163, 8 pages. http://dx.doi.

org/10.4102/abc.v44i1.163

Licensee: AOSIS

OpenJournals. This work is

licensed under the Creative

Commons Attribution License.

Read online:

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falz zhi code with your

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lb 2% to read online.

Province, South Africa

Background: Field studies confirmed that unusually long-tubed populations of Pelargonium

dipetalum from between Swellendam and Albertinia, Western Cape Province, South Africa,

are a distinct ecotype adapted to pollination by the long-proboscid fly, Prosoeca longipennis.

The geographical and morphological isolation of these populations suggests that they are

reproductively isolated from short-tubed populations, which are pollinated by bees.

Objectives: To determine and describe the floral variation in P. dipetalum, with a view to

recognising the long-tubed populations at some taxonomic level.

Method: All available collections were measured and compared.

Results: Populations of P. dipetalum were segregated into a short-tubed form with hypanthium

3 mm — 24 mm long and mostly pink petals that occurs from Betty’s Bay to Knysna, and a

long-tubed form with the hypanthium 34 mm — 54 mm long and consistently white petals

that is restricted to a small area east of Swellendam between Suurbraak and Albertinia. We

described the long-tubed form as the new subspecies P. dipetalum subsp. stenosiphon.

Conclusion: The new subspecies increases our understanding of the diversity in P. dipetalum

and represents a new taxon of conservation concern.

Introduction

The genus Pelargonium L’Hér. ex Aiton (Geraniaceae) includes approximately 280 species

distributed amongst 16 sections (Bakker ef al. 2004). Section Hoarea (Sweet) DC., with + 85 species

occurring mainly in the winter rainfall region of South Africa (Marais 2014), is one of the largest

sections in the genus. It comprises deciduous geophytes with carrot-shaped or turnip-shaped

tubers covered with papery bark and a highly condensed stem with the leaves in a radical tuft and

often dry at flowering (Marais 1994, 2014).

The section includes seven species with only the posterior two petals present, as opposed to the

full complement of five petals commonly found in the genus (Manning & Goldblatt 2012; Marais

1994). Pelargonium dipetalum L’Her. is the only member of this alliance occurring along the southern

coast in Western Cape Province and extends from Betty’s Bay to Knysna. The hypanthium in

P. dipetalum typically varies between 7 mm and 18 mm in length, but the species also includes a few

populations east of Swellendam with exceptionally long hypanthia, 35 mm — 50 mm long. These

populations were not recognised as a separate taxonomic entity by Marais (1994), who made no

correlation between their distribution and their distinctly longer hypanthium. This variant was,

however, identified by Manning and Goldblatt (2005) as representing a distinct morph adapted to

pollination by the long-proboscid fly Prosoeca longipennis Loew (Nemestrinidae).

A recent study of the P. longipennis pollination syndrome by Newman, Manning and Anderson

(2014) defined the guild (Fenster et al. 2004) as including mainly autumn-flowering species with

unscented, white to pink flowers with long floral tubes. A population of the long-tubed morph

of P. dipetalum included in the study confirmed that it is adapted to pollination by P. longipennis,

whereas the short-tubed morph of the species is visited by bees.

We document the distribution and variation in hypanthium length and petal colour in P. dipetalum

and confirm that the long-tubed populations represent a variant that is morphologically and

geographically discontinuous from the typical populations. We recognise these populations as

the new subspecies P. dipetalum subsp. stenosiphon.

http://www.abcjournal.org | doi:10.4102/abe.v44i1.163 ——

“ou nnn, PF ge 20 f 8 o 6) rj gi nal Re searc h SS

Research method and design

Fieldwork was conducted in the Langeberg area (Western

Cape Province, South Africa) and herbarium specimens from

the Bolus Herbarium (BOL), Compton Herbarium (NBG)

(including the South African Museum [SAM]) and the Pretoria

National Herbarium (PRE) were studied (abbreviations

according to Holmgren, Holmgren & Barnett 1990).

Hypanthium length and petal colour were recorded for all

specimens (Appendix 1). The shortest and longest values

for hypanthium length were recorded for each herbarium

collection, as well as the mean hypanthium length for all

short-tubed and long-tubed collections, respectively.

The distribution map was compiled using the quarter degree

reference system (Leistner & Morris 1976).

Results

Populations of P. dipetalum are separable into two groups

based on hypanthium length and petal colour. The common

form corresponding with the type of the species has short-

tubed flowers, with the hypanthium 3 mm — 23 mm long

(9.7 mm + 3.1 mm s.d.; n = 144) (Appendix 1; Figure 1),

mostly pink petals (rarely white in populations near Knysna

and Bredasdorp) (Figure 2) and a widespread distribution

along the coast from Betty’s Bay to Knysna (Figure 3).

Three populations from slightly further inland between

Suurbraak and Albertinia have consistently white petals

and a significantly longer hypanthium 34 mm — 54 mm long

(41.1 mm + 9.0 mm s.d.; n = 8) (Figures 1-3; Appendix 1).

There is no overlap in hypanthium length between the two

variants.

Length of flower tube (mm)

(o)

Pelargonium dipetalum

subsp. stenosiphon

Pelargonium dipetalum

subsp. dipetalum

FIGURE 1: Hypanthium length in the two subspecies of Pelargonium dipetalum,

showing the shortest and longest values plus the range in average shortest to

average longest values per herbarium collection.

Source: (a) Photographed by A. Johns; (b) photographed by J. Manning

FIGURE 2: Photographs of Pelargonium dipetalum showing flowers of the two subspecies, (a) Pelargonium dipetalum subsp. dipetalum from Kogelberg with pink flowers

and short hypanthium and (b) Pelargonium dipetalum subsp. stenosiphon from Albertinia with white flowers and long hypanthium.

——— http://www.abcjournal.org Pal doi:10.4102/abe.v44i1.163 wi

metres

| 300

600

metres

| --

Scale bar, 10 mm.

FIGURE 3: Geographical distribution of hypanthium length in Pelargonium dipetalum, showing the shortest (white bar) and longest (black bar) hypanthium length

recorded for each quarter degree grid, with (a) depicting, Pelargonium dipetalum subsp. dipetalum and (b) depicting Pelargonium dipetalum subsp. stenosiphon.

http://www.abcjournal.org | doi:10.4102/abc.v44i1.163

The two variants are geographically separated and are

morphologically distinct in hypanthium length and partially

in petal colour, but we were unable to find any other

differences between them. We accordingly treated the long-

tubed populations as comprising a separate subspecies

representing a distinct ecotype adapted to pollination by the

long-proboscid fly P. longipennis.

Taxonomic treatment

Pelargonium dipetalum L’Heér., Geranologia, seu Erodii,

Pelargonii, Monsoniae et Grieli historia iconibes illustrata:

t. 43 (1792). Geranium dipetalum (L’Her.) Poir.: 744 (1812).

Geraniospermum dipetalum (L’Heér.) Kuntze: 95 (1891).

Seymouria l’heritiert Sweet: 77 (1824-1826), nom. illegit. superfl.

pro P. dipetalum L’Her. Pelargonium Uhéritiert (Sweet) Don: 731

(1831). Type: Illustration in L’Heritier: t. 43 (1792) (icono.!).

Hoarea erythrophylla Eckl. & Zeyh.: 60 (1835). Pelargonium

erythrophyllum (Eckl. & Zeyh.) Steud.: 285 (1841). Type:

South Africa, [Western Cape], ‘Rivier Zonder Einde apud

villam Knoblauch (Zwellendam)’, Ecklon & Zeyher 457

[S, lecto., designated by Marais: 245 (1994)—JSTOR image!;

S, isolecto.—JSTOR image!] (JSTOR 2011a, 2011b).

Pelargonium niventt Harv.: 271 (1860). Geraniospermum nivenit

(Harv.) Kuntze: 95 (1891). Type: South Africa, [Western Cape],

‘elevated places in Sweetmilk Valley [Soetmelksvallei]’, Niven

s.n. [S, holo.—JSTOR image!] (JSTOR 2012).

Description

Geophytic herb up to 350 mm; tuber round to elongate,

sometimes with a moniliform root. Stipules: subulate and

adnate to petioles for two thirds of their length, 12 mm —

37 mm x 1 mm-—2 mm, ciliate. Leaves: erect; petioles 30 mm —

200 mm long, rigid, hirsute with appressed or patent hairs;

blades green or occasionally purple beneath, simple to

irregularly pinnatisect to bipinnatisect, elliptic in outline,

20 mm— 120mm = 13 mm—30 mm, acuminate, bases cuneate,

lamina hirsute above and appressed-hirsute beneath.

Inflorescence: 30 mm —200 mm long, scape with 2—3(5) pseudo-

umbels, each with (3)4-8(12) flowers; peduncles 25 mm —

150 mm long, with glandular hairs interspersed with bristle-

like hairs; bracts subulate, 3 mm — 7 mm = 1 mm — 2 mm,

hirsute beneath with distally appressed hairs; pedicels c.

0.5 mm long. Hypanthium: 3 mm — 54 mm long, reddish-

brown, densely covered with glandular hairs interspersed

with bristle-like hairs. Sepals: 5, lanceolate, acute, 5.0 mm —

11.5 mm x 1.2 mm — 4.0 mm, reflexed, green to reddish-

brown, indumentum as for peduncle. Petals: 2 in posterior

position, pink to white, usually with darker-coloured

nectar guides, obovate to spathulate, 12 mm — 19 mm

x 3 mm — 7(—-10) mm, emarginate, bases cuneate, recurved

BOX 1: Key to subspecies of Pelargonium dipetalum.

during anthesis. Stamens: white to pale pink, papillate; fertile

stamens 5, posterior filament shortest, 6.5 mm — 11.0 mm

long, lateral and anterior filaments 7.5 mm — 12.0 mm

long, free filaments wine-red; staminodes 2.0 mm — 5.5 mm

long; anthers dark red, 1.5 mm — 2.5 mm long, pollen orange.

Gynoecium: pink; ovary 3 mm — 6 mm long; style 1.5 mm —

6.0 mm long, lengthening during anthesis; stigma branches

1 mm — 3 mm long, wine-red. Fruit: bases of mericarps

5 mm — 8 mm long, eglandular, tails 20 mm — 31 mm long.

Description based on Marais (1994, 2000).

Pelargonium dipetalum subsp. dipetalum

Diagnosis

Petals pale to dark pink, rarely white; hypanthium

3 mm — 17(—23) mm long (Figure 2a).

Distribution

Subsp. dipetalum occurs from Betty’s Bay eastwards to

Belvedere near Knysna (Figure 3a).

Additional specimens seen

WESTERN CAPE.—3418 (Simonstown): Kogelberg Forest

Reserve above road to Oudebosch farm, (-BD), 30 Mar.

1971, Boucher 1457 (NBG); Kogelberg State Forest, halfway

between first turn and first bridge, left of road to Oudebos,

(-BD), 02 Mar. 1992, Kruger 407 (NBG, PRE); Fairy Glen,

firebreak, (-BD), 12 Mar. 1970, Boucher 1186 (NBG); Hangklip,

Betty’s Bay, north-east corner of plot 3009 near boundary

with school plot 2759, (-BD), 04 Mar. 1999, Forrester 1310

(PRE); Betty’s Bay, (-BD), 05 Feb. 1963, Levyns 11417 (BOL);

Betty’s Bay, lower mountain slopes above Anna’s Hof,

(-BD), 21 Feb. 1989, Rourke 1924 (NBG). 3419 (Caledon):

Central koppie, Haasvlakte, Houhoek, (-AA), 29 Jan. 1988,

Boucher & Stindt 5373 (NBG); Houhoek, Houtech terrain,

(-AA), 13 Apr. 1988, Boucher & Stindt 5395 (NBG); Houhoek,

Houtech terrain, (-AA), 16 Apr. 1988, Boucher & Stindt

5396 (NBG); Elgin, Caledon, (-AA), 24 Apr. 1943, Compton

14523 (NBG); Houhoek, at hotel, (-AA), 04 Apr. 1892,

Guthrie 2225 (NBG); near Palmiet River, Oudeburg, (-AA),

26 Apr. 1943, Leighton 503 (PRE); Houhoek, (-AA), 12 Apr.

1896, Schlechter 667 (PRE); Houhoek, (-AA), 12 Apr. 1896,

Schlechter 7561 (BOL); Elgin, at the Bridge, Caledon, (-AA),

06 Mar. 1926, Smith 2539 (PRE); Caledon, hill north of Baths,

(-AB), 31 Mar. 1922, Marloth 11085 (NBG, PRE); Kleinmond,

kloof north-west of Heuningklip, (-AC), 28 Apr. 1948, De

Vos 941 (NBG); Driebosch, Kleinmond, Caledon, (-AC),

01 Mar. 1943, Marais s.n. (NBG); Suikerboskop east of Bot

River Lagoon, (-AC), 25 Apr. 1975, Oliver 5798 (NBG, PRE);

Kleinmond, (-AC), 22 Apr. 1922, Stokoe 1327 (BOL); Groot

Hagelkraal near Pearly Beach, (-AC), 16 Mar. 1983, Van Wyk

1154 (NBG); Happy Valley, Riviersonderend Mountains,

(-BA), 12 Apr. 1941, Compton 10665 (NBG); flats at foot of

1A. Hypanthium 3 mm — 17(—23) mm long; petals pink, rarely Whit@.............ccceeeseeseeeeeee

1B. Hypanthium 35 mm —54 mm long; petals White...........cccccsecesecesseessccessetsestseeeseesseeeee

acu Rebuecue seb euk ouredd dior wieb a cuirak wOsu st aa lca MEK ca cLeu cRNA Ri cuestatnnuauc eae P. dipetalum subsp. dipetalum

Wicveconacded dss ccnte ss aoasberascvouchtcsvacestUanenscre ssodeeonUEMra Cen ecR onseonN P. dipetalum subsp. stenosiphon

http://www.abcjournal.org

{01:10.4102/abe.v44i1.163, —————

Riviersonderend Mountains near Neethlings Farm, (-BA), 29

Apr. 1950, Wilman 978 (PRE); Riviersonderend Mountains,

(-BB), Apr. to May 1950 [without day], Lewis SAM63209

(PRE); Zonderend near Olifantsbos, (-BB), 01 Apr. 1922,

Marloth 11090 (PRE); Riviersonderend, (-BB), 30 Apr.

1950, Middlemost 1509 (NBG); foothills of Riviersonderend

Mountains, (-BB), Apr. to May 1950 [without day], Stokoe

63208 (PRE); Sandies Glen near Sandfontein, north-east

sandstone slope of Koueberg, (-BC), 18 Mar. 1977, Hugo 908

(NBG, PRE); 4 km north-west of Napier, near Quarry Kop,

(-BD), 31 May 1995, Dreyer 525 (PRE); Skoenmakers River,

+ 10 km west of Napier, (-BD), 18 Mar. 1977, Thompson

3487 (NBG); Koks River, near entrance to farm, (-DA), 11

Mar. 1979, Hugo 1606 (NBG); Groot Hagelkraal, Hagelkraal

River area, north-east of farmstead, (-DA), 18 Apr. 1975,

Oliver 5895 (NBG); beacon 147, hills just north-west of Elim,

(-DA), 29 Mar. 1971, Oliver 3344 (NBG); Hagelkraal, poort

near Hagelkraal River, (-DA), 10 Mar. 1979, Thompson 3917

(NBG); on road from Elim to ‘Die Dam’, (-DA), 22 Mar.

1982, Van Wyk 801 (NBG); flats north-west of Soetanysberg,

Rietfontein Private Nature Reserve, (-DB), 25 Feb. 1994,

Beyers 224 (NBG); Bredasdorp, Rietfontein, (-DB), 13 Apr.

1978, Esterhuysen 34910 (BOL); near Mierkraal, south-west

of Bredasdorp, (-DB), 14 Mar. 1977, Hugo 823 (NBG);

4 miles [6.44 km] west of Elim, Bredasdorp, (-DB), 18 Feb.

1951, Maguire 845 (NBG); Vogel Valley, (-DB), 23 Apr. 1897,

Schlechter 1867 (PRE); Koks River, north-west of Buffeljagts

Mountain, (-DC), 17 Mar. 1978, Hugo 1197 (NBG). 3420

(Bredasdorp): De Hoop Nature Reserve, flats north of

station, (-AD), 17 Mar. 1977, Hugo 881 (NBG); De Hoop

Farm, (-AD), 1971 [without day or month], Van der Merwe

2015 (NBG); De Hoop, Potberg Nature Reserve, (-BC),

16 Feb. 1979, Burgers 1810 (NBG); De Hoop, Hamerkop,

(-BC), 29 Jan. 1985, Fellingham 869 (NBG); Zoetendal’s vlei,

east of Soetanysberg, (-CA), 25 Mar. 1982, Fellingham 398

(NBG), Uyshoek, 7 km due north of Arniston, (-CA), 15

Mar. 1977, Hugo 840 (NBG, PRE); Eiland’s Valley between

Bredasdorp and Skipskop, (-CA), 16 Mar. 1978, Hugo 1174

(NBG); hill near Bredasdorp, (-CA), [without date], Leighton

NBG 671/33 (BOL); De Hoop, Moerasfontein at second gate

from main road, (-CA), 31 Jan. 1985, Van Wyk 2171 (NBG);

De Hoop, Buffelsfontein, along road to Ryspunt, (-CB), 18

Mar. 1985, Van Wyk 2208 (PRE); Uitvlugt, south slopes of

Bredasdorp, (-DA), 16 Mar. 1978, Boucher 3775 (NBG). 3421

(Riversdale): Still Bay ridge north of rifle range, (-AD), 15

Feb. 1979, Bohnen 5071 (NBG, PRE); Schoemanshoek in

Albertinia, the Fisheries Road, (-BA), 14 Mar. 1978, Boucher

3720 (NBG); Cauca se Laagte, south of Albertinia, (-BC), 20

Mar. 1975, Oliver 5717 (NBG, PRE); Albertinia, Gouriqua/

Ystervarkpunt, (-BC), 20 Jan. 1987, Willemse 65 (NBG).

3422 (Mossel Bay): Mossel Bay, near new town extension

at Golden Rendezvous Hotel, (-AA), 29 Mar. 1987, Vlok

1855 (NBG); Knysna, Belvidere, (-BB), Feb. 1921 [without

day], Duthrie 500 (BOL, PRE, SAM). 3423 (Knysna): Knysna,

(-AA), 20 Feb. 1955, Adamson D331 (PRE); Knysna, (-AA),

Mar. 1921 [without day], Breyer 25210 (PRE); Knysna, (-AA),

Feb. 1921 [without day], Breyer 23973 (PRE); Knysna Heads,

(-AA), Feb. 1922 [without day], Fourcade 2018 (BOL).

http://www.abcjournal.org ee doi:10.4102/abce.v44i1.163

® Original Research ——-——---——

Without precise locality: Caledon division, Purcell 45919

(SAM).

Pelargonium dipetalum subsp. stenosiphon

J.C.Manning & M.M.le Roux, subsp. nov.

Type: SOUTH AFRICA. Western Cape: (3421) Riversdale:

2 km west of Dekriet siding on main road, (-AB), 10 Apr.

1979, Bohnen 5551 (NBG, holo.).

Diagnosis

Petals white; hypanthium 35 mm — 54 mm long (Figure 2b).

Distribution

Subsp. stenosiphon is localised along the southern foothills of

the Langeberg west of Swellendam, between Suurbraak and

Albertinia (Figure 3b).

Conservation

Unlike subsp. dipetalum, which is relatively widespread and

partially protected in nature reserves, subsp. stenosiphon

has a much more limited range. The taxon is recorded from

an area 90 km x 20 km in extent, much of which is under

pressure from agricultural activity. No known populations

are formally protected and the conservation status of the

subspecies needs to be assessed.

Additional specimens seen

WESTERN CAPE.—3420 (Bredasdorp): Swellendam,

Zuurbraak [Suurbraak], (-BA), 01 Mar. 1930, Thode A2301

(NBG, PRE). 4321 (Riversdale): flats west of Dekriet siding

west of Albertinia, (-AB), 19 Mar. 1975, Oliver 5692 (NBG).

Discussion

The two subspecies of P. dipetalum are distinguished by

hypanthium length and mostly also by petal colour (Figure 1

and Figure 2). The flowers in subsp. dipetalum are usually

pale to dark pink, but two populations, one near Bredasdorp

and the other at Belvedere near Knysna, have white flowers,

and the hypanthium ranges from 3 mm to 23 mm long. The

petals in subsp. dipeatlum are always marked with dark

nectar guides. In contrast, the petals in subsp. sfenosiphon

are consistently white with only faint nectar guides and the

hypanthium is 35 mm — 54 mm long.

The floral differences between the two subspecies of

P. dipetalum are associated with a shift in pollination system

from bee-pollination in the typical ecotype to pollination by

the long-proboscid fly P. longipennis in subsp. stenosiphon.

In an analogous situation, Becker and Albers (2010) recently

described P. alternans subsp. longicalcar M.Becker & F.Albers

(2010) for populations from the Outeniqua Mountains in

the Little Karoo differing from the other two subspecies in

having a longer hypanthium [(12—)18 mm — 36 mm long

in P. alternans subsp. longicalcar versus 3 mm — 9 mm in

subsp. alternans and subsp. parviinflorescens M.Becker &

F.Albers (2010)]. The long-tubed subspecies in this species

is also thought to be pollinated by a long-proboscid fly

and has undergone evolutionary changes similar to those

documented for P. dipetalum subsp. stenosiphon.

Acknowledgments

Amida Johns kindly provided the image of P. dipetalum subsp.

dipetalum, taken in the Kogelberg Nature Reserve.

Competing interests

The authors declare that they have no financial or personal

relationships that may have inappropriately influenced them

in writing this article.

Authors’ contributions

M.M.1.R. (South African National Biodiversity Institute)

and J.C.M. (South African National Biodiversity Institute)

collectively performed the research. J.C.M. conducted the

fieldwork and provided input during the compilation of the

manuscript; M.M.1.R. compiled the manuscript.

References

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of Pelargonium (Geraniaceae) based on DNA sequences from three genomes’, Taxon

53, 17-28. http://dx.doi.org/10.2307/4135485

Becker, M. & Albers, F., 2010, ‘Pollinator shift and speciation in Pelargonium alternans

(Geraniaceae)’, Schumannia 6, 207-218.

Don, G., 1831, A general system of gardening and botany, vol. 1, Rivington, London.

Ecklon, C.F. & Zeyher, K.L.P., 1835, Enumeratio plantarum africae australis extra-

tropicae, vol. 1., Prostat apud Perthes & Besser, Hamburg.

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Fenster, C.B., Scott Armbruster, W.S., Wilson, P., Dudash, M.R. & Thompson, J.D.,

2004, ‘Pollination syndromes and floral specialization’, Annual Review of Ecology

Evolution and Systematics 35, 375-403. http://dx.doi.org/10.1146/annurev.

ecolsys.34.011802.132347

Harvey, W.H., 1860, ‘Geraniaceae’, in W.H. Harvey & O.W. Sonder (eds.), Flora capensis,

vol. 1, A.S. Robertson, Cape Town.

Holmgren, P.K., Holmgren, N.H. & Barnett, L. (eds.), 1990, Index herbariorum. Part 1:

The herbaria of the world, 8th edn., New York Botanical Garden, New York.

JSTOR, 2011a, Isolectotype of Hoarea erythrophylla Eckl. & Zeyh. [family Geraniaceae],

viewed 14 May 2014, from http://plants.jstor.org/specimen/s09-40774?s=t

JSTOR, 2011b, Lectotype of Hoarea erythrophylla Eckl. & Zeyh. [family Geraniaceae],

viewed 14 May 2014, from http://plants.jstor.org/specimen/s09-40679 ?s=t

JSTOR, 2012, Holotype of Pelargonium nivenii Harv. [family Geraniaceae], viewed

14 May 2014, from http://plants.jstor.org/specimen/s09-40660?s=t

Kuntze, C.E.O., 1891, Revisio generum plantarum, Arthur Felix, Leipzig.

UHeritier de Brutelle, C-L., 1792, Geranologia, seu Erodii, Pelargonii, Monsoniae et

Grieli historia iconibes illsutrata, Petri-Francisci Didot, Paris.

Leistner, O.A. & Morris, J.W., 1976, ‘Southern African place names’, Annals of the Cape

Provincial Museums 12.

Manning, J.C. & Goldblatt, P., 2005, ‘Radiation of pollination systems in the Cape

genus Tritoniopsis (|Iridaceae: Crocoidea) and the development of bimodal

pollination strategies’, International Journal of Plant Science 166, 459-474. http://

dx.doi.org/10.1086/428703

Manning, J.C. & Goldblatt, P., 2012, ‘Pelargonium saxatile (Geraniaceae: Section

Hoarea), a new species from the southwestern Cape, South Africa, and a key

to the species of the P. dipetalum group’, South African Journal of Botany 78,

266-269. http://dx.doi.org/10.1016/j.sajb.2011.04.007

Marais, E.M., 1994, ‘Taxonomic studies in Pelargonium section Hoarea (Geraniaceae)’,

PhD thesis, Department of Botany, University of Stellenbosch.

Marais, E.M., 2000, Taxonomic studies in Pelargonium section Hoarea (Geraniaceae),

The Geraniaceae Group, England. :

Marais, E.M., 2014, ‘One name change and three new species of Pelargonium,

section Hoarea (Geraniaceae) from the Western Cape Province’, South African

Journal of Botany 90, 118-127. http://dx.doi.org/10.1016/j.sajb.2013.10.013

Newman, E., Manning, J.C. & Anderson, B., 2014, ‘Matching floral and pollinator traits

through guild convergence and pollinator ecotype formation’, Annals of Botany

113, 373-384. http://dx.doi.org/10.1093/aob/mct203

Poiret, J.L.M., 1812, ‘Encyclopédie methodique’, in J.B-A.P.M. Lamarcke (ed.), Botanique,

Supplementum, vol. 2, p. 2, Panckouke, Paris.

Steudel, E.G., 1841, Nomenclator botanicus, 2nd edn., vol. 2., J.G. Cotta, Stuttgart.

Sweet, R., 1824-1826, Geraniaceae, vol. 3., James Ridgway, London.

Appendix starts on the next page >

/www.abcjournal.org 6 | doi:10.4102/abc.v44i1.163 —— - -

Page 7 of 8 Original Research

Appendix 1

APPENDIX 1: Hypanthium length in Pelargonium dipetalum subspp. dipetalum and stenosiphon. Measurements are for the shortest and longest hypanthium represented

on each herbarium collection studied.

Subspecies Collector(s) and number Herbarium Hypanthium length (mm)

Shortest Longest

Pelargonium dipetalum subsp. dipetalum Boucher 1487 NBG 12 17

Boucher 1186 NBG tat 11

Forrester 1310 PRE 8 8

Kruger 407 NBG 10 LS,

Kruger 407 PRE 2) 15

Levyns 11417 BOL 13 14

Rourke 1924 NBG 10 13

Boucher and Stindt 5373 NBG 11 12

Boucher and Stindt 5396 NBG 9 11

f Boucher and Stindt 5395 NBG 12 13

Compton 14523 NBG 10 12

Guthrie 2225 NBG 8 12

Leighton 503 PRE 9 13

Schlechter 667 PRE 10 14

Schlechter 7561 BOL 7 g

Smith 2539 PRE

Marloth 11085 NBG 7

Marloth 11085 PRE 6 7

De Vos 941 NBG 10 23

Marais s.n. NBG 9 11

Oliver 5798 NBG

Oliver 5798 PRE 5 8

Stokoe 1327 BOL 10 13

Van Wyk 1154 NBG 8 13

Compton 10665 NBG i) 10

Wilman 978 PRE 9 10

Lewis SAM 63209 PRE 7 10

Marloth 11090 PRE 7 14

Middlemost 1509 NBG 8 13

Stokoe 63208 PRE 7 13

Hugo 908 NBG 11 13

Hugo 908 PRE is) 16

Dreyer 525 PRE 7 9

Thompson 3487 NBG 10 15

Hugo 1606 NBG 10 15

Oliver 5895 NBG 6 11

Oliver 3344 NBG 8 11

Thompson 3917 NBG 7 10

Van Wyk 801 NBG 8 9

Beyers 224 NBG 8 10

Esterhuysen 34910 BOL 9 2.

Hugo 823 NBG 8 10

Maguire 845 NBG 7 12

Schlechter 1867 PRE 6 9

Hugo 1197 NBG tla 15

Hugo 881 NBG 7h 9

Van der Merwe 2015 NBG 9 bse

Burgers 1810 NBG 14 15

Fellingham 869 NBG 9 10

Fellingham 398 NBG 3 i

Hugo 840 NBG 10 14

Hugo 1174 NBG 4 )

Hugo 840 PRE 7 9

Leighton NBG 671/33 BOL 7 11

Van Wyk 2171 NBG 8 9

Van Wyk 2208 PRE 12 14

Boucher 3775 NBG 9 16

Bohnen 5071 PRE 4 5

Source: Herbaria names abbreviated according to Holmgren, P.K., Holmgren, N.H. & Barnett, L. (eds.), 1990, Index herbariorum. Part 1: The herbaria of the world, 8th edn., New York Botanical

Garden, New York

NBG, Compton Herbarium; PRE, the Pretoria National Herbarium; BOL, the Bolus Herbarium; SAM, the South African Museum.

Appendix 1 continues on the next page >

Sass a —— http://www.abcjournal.org doi:10.4102/abc.v44i1.163 ——— Sa SHESTE

APPENDIX 1 (Continues...): Hypanthium length in Pelargonium dipetalum subspp. dipetalum and stenosiphon. Measurements are for the shortest and longest hypanthium

represented on each herbarium collection studied.

Subspecies Collector(s) and number Herbarium Hypanthium length (mm)

Shortest Longest

Bohnen 5071 NBG 8 9

Boucher 3720 NBG 8 9

Oliver 5717 NBG 8 2)

Oliver 5717 PRE 8 9

Willemse 65 NBG 5 8

Vlok 1855 NBG 9 10

Duthrie 500 SAM 6 9

Duthrie 500 PRE 4 13

Duthrie 500 BOL 7 9

Adamson D331 PRE 8 9

Breyer 25210 PRE 10 13

Breyer 23973 PRE 6 )

Fourcade 2018 BOL 4 8

Purcell 45919 SAM 4 8

Pelargonium dipetalum subsp. Thode A2301 NBG 34 54

stenosiphon Thode A2301 PRE 34 34

Bohnen 5551 NBG 35 36

Oliver 5692 NBG 35 47

Source: Herbaria names abbreviated according to Holmgren, P.K., Holmgren, N.H. & Barnett, L. (eds.), 1990, Index herbariorum. Part 1: The herbaria of the world, 8th edn., New York Botanical

Garden, New York

NBG, Compton Herbarium; PRE, the Pretoria National Herbarium; BOL, the Bolus Herbarium; SAM, the South African Museum.

Sea aa ML OTAG) few Article ———

Factors influencing the adaptation and distribution of

Colophospermum mopane in southern Africa’s mopane

Authors:

Rudzani A. Makhado!

Isaac Mapaure?

Martin J. Potgieter*

Wilmien J. Luus-Powell*

Amani T. Saidi?

Affiliations:

1Department of Biodiversity,

University of Limpopo,

South Africa

?Research and Publications

Office, University of Namibia,

Namibia

South African Environmental

Observation Network,

National Research

Foundation, South Africa

Correspondence to:

Rudzani Makhado

Email:

makhado2002@yahoo.com

Postal address:

Private Bag X1106, Sovenga,

South Africa

Dates:

Received: 05 May 2014

Accepted: 31 July 2014

Published: 12 Nov. 2014

Republished: 13 Nov. 2014

How to cite this article:

Makhado, R.A., Mapaure, I.,

Potgieter, M.J., Luus-Powell,

W.J. & Saidi, A-T., 2014,

‘Factors influencing the

adaptation and distribution

of Colophospermum mopane

in southern Africa’s mopane

savannas — A review’,

Bothalia 44(1), Art. #152,

9 pages. http://dx.doi.

org/10.4102/abc.v44i1.152

Note:

This article was republished

with the correct R? values

reflecting in Figure 3.

Read online:

Scan this QR

code with your

smart phone or

mobile device

to read online.

savannas — A review

Colophospermum mopane is the dominant tree or shrub within mopane woodland in the

subtropical areas of southern Africa’s savanna ecosystems. This article provided a review

on the adaptation capabilities of mopane against fire, browsing activity and environmental

stresses. It further reviewed and tested the extent to which rainfall, temperature, altitude and

soil types had an effect on the distribution of mopane in southern Africa. Mopane is adapted

to survive moisture stresses, low nutrient environments and even disturbances caused

by fire and browsing by large herbivores through its physical, physiological and chemical

responses. Adaptation of mopane to various stresses enables it to dominate the low-lying

areas of southern Africa’s savannas. The distribution of mopane is best associated with low

to moderate rainfall (R* = 0.38), high temperature (R* = 0.42), low altitudes (R* = 0.44) and a

variety of soil types. An increase in the annual rainfall (> 800 mm) and altitude (> 800 m.a.s.1.),

coupled with a reduction in the minimum temperature and acidic soil, limits the distribution

of mopane. Mopane in South Africa occurs under similar environmental conditions to those

in Zimbabwe and Zambia, but quite different from those in Angola, Namibia, Mozambique,

Malawi and Botswana where mopane occurs.

Introduction

Colophospermum mopane (Kirk ex Benth.) Kirk ex J.Léonard, commonly known as mopane, is

the dominant tree or shrub within mopane woodland in the subtropical areas of southern

Africa’s savanna ecosystems, between latitudes 9° S and 25° S (Henning 1976; Mapaure 1994;

Sebego 1999; Werger & Coetzee 1978; White 1983). Estimates show that mopane woodland

accounts for about 30% — 35% of the 1.5 million km? of savannas in southern Africa (Mapaure

1994; White 1983), which represents more than a quarter of land area in the region. Mopane is

distributed in the hot, dry, valley bottoms and adjacent plains of southern Angola and northern

Namibia, across Botswana and Zimbabwe to central and southern Mozambique, and from the

Luangwa valley in Zambia and central Malawi to northern South Africa (Mapaure 1994; Porter

1968; Siebert 2012; Timberlake, Chidumayo & Sawadogo 2010; Werger & Coetzee 1978; White

1983) (Figure 1). The total area covered by mopane woodland in the whole of southern Africa

is 555 000 km? (Mapaure 1994) (Table 1).

Previous studies have demonstrated that rainfall, altitude and soil types influence the distribution

of mopane in southern Africa (e.g. Burke 2006; Cole 1986; Mapaure 1994; Voorthuizen 1976;

Werger & Coetzee 1978). Mopane occurs in areas receiving low to moderate annual rainfall

ranging from 400 mm to 800 mm (Madams 1990; Thompson 1960; Werger & Coetzee 1978).

These are normally areas at altitudes ranging from 200 m.a.s.l. to 600 m.a.s.l. (White 1983), with

variable soils, but usually fine-grained, having textures ranging from sandy through loamy to

clayey. The species is also known to occupy both shallow and deep soils, containing significant

amounts of exchangeable sodium (Madams 1990; Thompson 1960; Werger & Coetzee 1978).

Other factors influencing the distribution of mopane include minimum temperature and dry

season day length (Stevens et al. 2014). Mopane is commonly distributed in high temperature

areas (Table 2) and minimum temperature of < 5 °C limits its distribution (Burke 2006; Cole

1986; Henning 1976; Stevens et al. 2014; Timberlake, Nobanda & Mapaure 1993; Werger &

Coetzee 1978; White 1983; Whitecross, Archibald & Witkowski 2012), especially in its southern

range (Stevens et al. 2014). However, although mopane is predominantly in frost free areas, the

species is capable of withstanding light frost (Thompson 1960) and tall mopane trees of > 4m

in height can survive minimal frost damage (Whitecross et al. 2012).

http://www.abcjournal.org/ | 9 | dod O41 02/abewA4 ile 2 SSS

Mopane is considered an important plant species to

people, and wild and domestic animals in its distribution

in southern Africa. Rural dwellers use it for firewood

(Liengme 1983),

(Makhado et al. 2009) and, to a lesser extent, for medicinal

purposes (Madzibane & Potgieter 1999; Mashabane, Wessels

& Potgieter 2001). In some parts of the region, there has

been increasing use of mopane in urban areas for firewood

construction of traditional structures

as the cost of electricity keeps increasing. Mopane also

hosts mopane worms, larvae of the moth Imbrasia belina,

which are consumed for their nutritional value (Dreyer &

Wehmeyer 1982; Voorthuizen 1976) and traded to generate

income (Styles 1996). Dry mopane leaves, twigs and pods

provide a valuable source of browse for wild animals such as

elephants (Ben-Shahar & MacDonald 2002) and greater kudu

* y ”

— ct in te

\ [3

c Y ¢ (4

\ x . — >» y

\ 5

) i

X | —_ “7

a ola Pe

f Aig Zambia AZ

{ IF £ oa

f Ye Ze

(ee eeu

\ pu bs J = 4 pee a

\ Namibia_ Botswana oy TaN

b 4 NK < eS

{ Sf Se / ;

) \ ae aches: ; ~ i ‘

| \ —South Africa 5

| : I a = J

J \ \ A re Ve

\ . YA

| N J wa a=

Be | _} Country boundaries

___} Mopane woodland distribution

6 2000 Kilometres

Source: This map is an extract from Mucina and Rutherford’s (2006) data on the vegetation

of South Africa, Lesotho and Swaziland (VegMap) and White’s (1983) data on vegetation

of Africa. For more information, please consult the reference list of the article: Makhado,

R.A., Mapaure, |., Potgieter, M.J., Luus-Powell, W.J. & Saidi, A.T., 2014, ‘Factors influencing

the adaptation and distribution of Colophospermum mopane in southern Africa’s mopane

savannas — A review’, Bothalia 44(1), Art. #152, 9 pages. http://dx.doi.org/10.4102/abc.

v44i1.152

FIGURE 1: The distribution of mopane-dominated woodlands in southern

Africa.

Review Article ———

(Hooimeijer et al. 2005), especially during the dry season

and drought periods (Bonsma 1942; Macala, Sebolai &

Majinda 1992; Mosimanyana & Kiflewahid 1988; Timberlake

1995), when the tannins have leached out. In addition, the

secretion of Arytaina mopane nymphs, commonly known as

lerp, increases the palatability of mopane leaves (Ross 1977;

Van Wyk 1972), because the lerps have high sucrose content

(Styles 1993). The lerps are highly sought after by baboons,

monkeys, birds (Herremans-Tonnoeyr & Herremans 1995)

and even humans, especially in the northern part of South

Africa (Pettey 1925) and Botswana (Sekhwela 1989), because

of their sweetness; they contain about 53% water-soluble

sugars (Sekhwela 1989).

Considering the extensive distribution of mopane in the

low-lying areas of southern Africa and its importance to

human livelihoods, domestic and wild animals throughout

its distribution range, it becomes a research challenge

when factors influencing its distribution are not easily

detectable (Siebert 2012) or not even well understood

(Stevens ef al. 2014). Various sources have contributed to

the understanding of mopane distribution (e.g. Cole 1986;

Du Plessis 2001; Henning 1976; Madams 1990; Mapaure 1994;

Thompson 1960; Timberlake 1995; Timberlake ef al. 1993;

Werger & Coetzee 1978; White 1983), but there is still a gap

in identifying the underlying factors influencing mopane

distribution in southern Africa (Siebert 2012; Stevens

et al. 2014). This is creating a subsequent gap in our ability to

TABLE 1: Area covered by mopane-dominated woodland in southern Africa.

Country Area (km?) Proportion of Proportion of total

country area(%) mopane area (%)

Angola 112 500 9 20

Zimbabwe 101 500 26 18

Mozambique 98 000 13 18

Botswana 85 000 aS 15

Namibia 77 000 9 14

Zambia 43 500

South Africa 23 000 2

Malawi 10 000 9 2

Total 550 500 89 100

Source: These data are taken from Mapaure, |., 1994, ‘The distribution of Colophospermum

mopane (Leguminosae-Caesalpinioideae) in Africa’, Kirkia 15, 1-5

TABLE 2: Environmental factors associated with the distribution of mopane in southern Africa.

Country Mean annual rainfall (mm) Mean daily temperature (°C) _—_Alltitudinal range (m.a-s.1.) Soil types References

Angola 100-600 16-25 100-1200 g,S,a S727 AS)

Zimbabwe 450-700 16-30 300-950 c, so, |, g, ss, fe il (3), 7/, 8)

Mozambique 400-700 20-29 100-500 c, Is, s, bs, a 1,2,6,9

Botswana 400-600 13-30 800-900 s, si, cl, c, b, h tL, 2, 3, 8)

Namibia 50-600 12-31 150-1000 Lf 216) FS; Oy dd

Zambia 600—1000 14-30 400-1000 a, fy, sl, g, |, ss, fe, h il 2, 5), 6, 7,2)

South Africa 250-650 15-31 200-800 a,g,b,c,s,| 1, 3,4, 6,9, 10

Malawi 700-800 19-28 450-500 sc, m 1259

Minimum 50-550 12-25 100-500 - >

Maximum 700—1000 20-31 800-1200 =

Average 369-700 16-29 313-856 5 =

Sources: 1, Mapaure (1994); 2, Werger and Coetzee (1978); 3, Acocks (1953); 4, Mucina and Rutherford (2006); 5, Porter (1968); 6, Henning (1976); 7, Madams (1990); 8, Erkkila and Siiskonen

(1992); 9, Du Plessis (2001); 10, Rutherford et a/. (200

6); 11, Okitsu (2005).

For more information, please consult the reference list of the article: Makhado, R.A., Mapaure, |., Potgieter, M.J., Luus-Powell, W.J. & Saidi, A.T., 2014, ‘Factors influencing the adaptation and

distribution of Colophospermum mopane in southern Africa’s mopane savannas — A review’, Bothalia 44(1), Art. #152, 9 pages. http://dx.doi.org/10.4102/abce.v44i1.152

Soil types symbols: a, alluvial; b, basaltic; bs, brown soils; c, clayey; cl, clay loamy; f, ferruginous; fe, ferrallitic; fv, fluvisol-vertisol; g, granitic; h, halomorphic; |, lithosols; Is, loamy sand; m,

mopanosols; s, sandy; sc, sandy clays; sl, sandy-loamy,

; Si, silt; so, sodic; ss, sandstone.

http //

Iwww.abcjournal.org/

doi:10.4102/abc.v44i1.152 ——————

effectively manage mopane and the wild animals it supports.

Current climatic changes provide further complexities for

predicting the distribution of mopane. As a result, there

is a need to adequately review existing information in an

integrated manner. This will allow a better understanding of

the factors influencing the distribution of mopane in southern

African. This review is also critically important because it

gives insight into the potential future distribution scenarios of

mopane. An extensive review of a mixture of literature (i.e.

journal articles, books, conference proceedings and reports),

specifically dealing with adaptation of mopane and factors

influencing its distribution in southern Africa was carried

out. The aim of this article is therefore to provide a review of

the mechanisms that enable mopane to survive disturbances

caused by fire, browsing activity by large herbivores and

environmental stresses in the savanna ecosystem. The

article further tested the effect of environmental factors on

the distribution of mopane in the southern Africa’s savanna

ecosystem.

Adaptation of mopane to fire and

browsing activity

Mopane is widely distributed in the southern Africa’s

savanna (Mapaure 1994), an ecosystem which supports

frequent fires (Andreae et al. 1994; Kennedy 2000; Scholes

1995) and large herbivores (Sankaran & Anderson 2009). Fire

negatively affects the morphology of mopane (Gandiwa &

Kativu 2009; Mlambo & Mapaure 2006), destroys the aerial

components of mopane shrubs (Henning 1976) and causes

a reduction in mopane height and stem circumference

(Kennedy & Potgieter 2003).

In addition, mopane is highly browsed by large herbivores

such as elephants (Ben-Shahar 1993; Ben-Shahar &

MacDonald 2002; Smallie & O’Connor 2000), mainly owing

to its high nutritional value (Ben-Shahar & MacDonald 2002;

Bonsma 1942; Macala et al. 1992; Mosimanyana & Kiflewahid

1988). Elephants’ preference for mopane makes it susceptible

to elephant-induced damage (Lewis 1991). Elephants’

feeding behaviour can transform mopane woodlands to

coppiced shrubby stands. Furthermore, elephants also

inhibit height recruitment of mopane by repeatedly breaking

the branches, ring-barking, heavy browsing and toppling

the tree (Lewis 1991; Smallie & O’Connor 2000). As a result,

fire and browsing activity has a notable effect on mopane

structure, which also has implications on the growth and

distribution of mopane in southern Africa’s savannas.

Despite the disturbances caused by fires and browsing

activity by large herbivores, mopane is capable of surviving

through its coppicing ability and production of chemicals for

defence. Various authors have shown that mopane coppices

rapidly (Luoga, Witkowski & Balkwill 2004; Mlambo

& Mapaure 2006; Mushove 1992; Mushove & Makoni

1993; Tietema 1989) after it has been disturbed by fire and

browsing animals. In addition, mopane wood contains

crystals of calcium oxalate, which contribute to high wood

density (Prior & Cutler 1992) and also enhance resistance of

SaaS http://www.abcjournal.org/

Review Article

the wood to fire (Centro Informatico Cientifico de Andalucia

[CICA] 1996). These crystals effect the burning properties

of the wood through producing considerable amounts of

carbon dioxide, which retards the fire flame (CICA 1996).

During the growing season, mopane also produces a high

concentration of secondary metabolites, such as tannins

and phenols, in order to deter herbivores from browsing it

(Kohi et al. 2010; Wessels, Van der Waal & De Boer 2007),

regardless of its high nutritional value. Therefore, the

ability of mopane to coppice after disturbances and produce

chemical defence enables it to survive disturbances caused

by fire and browsing animals.

Adaptation of mopane to

environmental stresses

It is well documented that mopane has the ability to survive

low to moderate rainfall (Henning 1976; Timberlake 1995),

water stresses (Choinski & Tuohy 1991; Mantlana 2002) and

high temperatures (Dye & Walker 1980; Henning 1976), but

how the species is able to survive such ‘harsh’ environmental

conditions in southern Africa has not been adequately

reviewed. It is the physical characteristics (e.g. Henning

1976; Madams 1990) and physiological mechanisms (e.g.

Choinski & Tuohy 1991; Dye & Walker 1980; Henning 1976;

Johnson et al. 1996; Mantlana 2002), which enable mopane to

tolerate water stress and high temperature conditions. It is

the response of mopane roots and leaves to changes in the

surrounding environment that enables it to survive these

‘harsh’ environmental conditions and dominate most low-

lying parts of southern Africa’s savanna. The physical and

physiological abilities of mopane are therefore discussed in

this article in order to better explain the mechanisms that

enable the species to survive in hot, dry, low-lying areas of

southern A frica.

Root-related adaptations

Mopane is essentially a shallow-rooted species (Henning

1976; Smit & Rethman 1998) and has high root biomass

(Smit & Rethman 1998). It is considered a shallow-rooted

species because its roots are mainly found at a depth of

20 cm — 120 cm (Mantlana 2002; Smit & Rethman 1998;

Thompson 1960), but can also reach 2 m in deep soils

(Mantlana 2002; Sebego 1999; Timberlake & Calvert 1993).

Mantlana (2002) indicated that the total root density for

short and tall mopane was highest in the first 20 cm of the

soil profile and then declined with increase in soil depth.

The combination of a shallow rooting system and high

root biomass places mopane in a competitive advantage

in areas where conditions lead to the development of a

zone of maximum water retention and nitrogen near the

surface (Dye & Walker 1980; Henning 1976; Mlambo,

Nyathi & Mapaure 2005; Smit & Rethman 1998). The

high, fine root densities of mopane, especially at a depth

of 20 cm — 120 cm (Mantlana 2002; Smit & Rethman 1998;

Thompson 1960), are important as they facilitate quick

water and nutrient acquisition and transport (Madams

1990; Mantlana 2002).

SCO OA 100) ADC NAA INA SDN ae

Another advantage is that the B horizon under mopane sodic

soils is relatively impermeable (Dye & Walker 1980), which

provides more moisture retention to the A and O Horizons

where most of mopane roots are found. The relatively

impermeable B horizon further restricts moisture from

filtrating down to the C horizon. As indicated by Dye and

Walker (1980), these characteristics enable shallow-rooted

species such as mopane to have a competitive advantage for

moisture uptake over deep-rooted species. It is believed that

the shallow rooting system of mopane, complemented by

its high root biomass, enables it to quickly absorb and store

the available moisture and nutrients near the soil surface,

enabling it to survive the ‘harsh’ environmental conditions

of southern Africa’s savanna.

The roots play a further critical role in the survival of

mopane. Mopane coppices easily (Mushove 1992; Mushove

& Makoni 1993; Tietema 1989; Tietema, Kgathi & Merkesdal

1988), mainly because its roots have the ability to produce

root suckers, which enables the shoots to grow faster than

newly established seedlings (Luoga et al. 2004). As indicated

by Mantlana (2002), the ability of mopane roots to coppice

confers a degree of resilience to natural and anthropogenic

disturbance, which is critical in ensuring its survival.

Cell sap-related adaptations

Stressed mopane shows a marked increase in relative

nitrogen content, which suggests that the resistance of

mopane to severe soil moisture stress is partly caused by the

build-up of soluble nitrogenous compounds within the cell

sap. In addition, the uptake of magnesium also plays a direct

role in the maintenance of water use efficiency of mopane

by catalysing the metabolic production of organic solutes,

thereby increasing the osmotic pressure of the cell sap and

thus enhancing the ability of mopane to withstand moisture

stresses (Henning 1976).

Leaf-related adaptations

Mopane is physiologically adapted to dry (moisture-

stressed) environmental conditions (e.g. Choinski & Tuohy

1991; Dye & Walker 1980; Prior 1991). It is adapted through

restricting transpiration, a mechanism that enables the

species to maintain high water potential (Henning 1976).

This is largely through folding the leaves, stomatal responses

and osmotic adjustment, which are considered critical

mechanisms in enabling mopane to survive water-stressed

and high temperature conditions of southern Africa.

Leaf responses

The mopane leaf has two triangular leaflets shaped like wings

of a butterfly. The leaves are leathery and resinous (Henning

1976). The leathery membrane on the leaf acts as a buffer

layer to avoid direct heat from the sun and also reduces the

rate of water loss through evapotranspiration. Mopane also

has a tendency to fold its leaflets together, especially during

the heat of the day (Madams 1990; Timberlake 1995). When

the leaflets are folded together, especially when the leaf

http://www.abcjournal.org/

ReviewACle(S=—S

temperature exceeds 30 °C, it reduces direct heat from the

sun. This means that the few exposed stomata will close,

which also assists in reducing the loss of water through

evapotranspiration. When the stomata are closed, the rate

of photosynthesis is reduced, confirming the findings by

Prior (1991), who indicated that photosynthesis by mopane

leaves is lower during the heat of the day than during the

night or cool days when the temperature is relatively low. In

addition, mopane is a deciduous species and sheds its leaves

during the dry season, mainly from August to October. The

ability of mopane to lose its leaves enables it to conserve

water that could have been lost during evapotranspiration.

Stomatal responses

Stomata are randomly distributed on the adaxial (top) and

abaxial (bottom) surface on mopane leaves, occupying

mainly the minor veins (Potgieter & Wessels 1998). However,

mopane has fewer stomata on the exposed adaxial leaf

surface compared with most other species (Prior 1991), which

implies that the stomata are mainly distributed on the abaxial

surface. The few exposed stomata on the adaxial surface of

mopane leaves therefore limit the number of openings on

the leaf; hence less moisture is lost through the leaves. It has

also been shown that mopane stomatal conductance declines

almost linearly at light saturation from March to August, at

585mmol m°*s!— 172mmol m°s', respectively (Mantlana

2002). The decline in mopane stomatal conductance occurs

when the soil moisture is low, especially during dry seasons

and drought conditions. This mechanism enables mopane

to conserve water during hot, dry conditions. Mantlana

(2002) suggested that the reduction in stomatal conductance

observed when soil water deficit increased may be explained

by the reduction in predawn leaf water potential. However,

this article states that leaf temperature needs to be taken into

account because it also has an influence on mopane stomatal

conductance. Nevertheless, the ability of mopane to close the

stomata during high temperature and water-stress periods

enables it to reduce the loss of moisture and nutrients, which

is critical in ensuring its future survival under the hot, dry

conditions of southern Africa.

Osmotic adjustment

The ability of mopane to grow and tolerate water-stressed

conditions is also through its osmotic adjustment (Henning

1976; Timberlake 1995). Osmotic adjustment processes lower

cell osmotic potential, thereby enabling intercellular water to

flow towards the inside of cells. This process is an important

mechanism in maintaining cell turgor pressure under

reduced soil water potential and thus enables the plant to

tolerate drought or water-limiting conditions (Chen & Jianga

2010; Hsiao et al. 1976).

As a result of the osmotic adjustment, mopane has the

ability to germinate and establish root growth at lower

water potentials than otherwise would be possible. The

seeds of mopane can germinate and withstand water stress

from -0.2 MPa to -0.51 MPa without wilting (Choinski &

Tuohy 1991; Henning 1976; Johnson ef al. 1996). Although

alo lO eNO el eu NS 2 ee

Page 5 of 9

the predawn xylem pressure potential analysis for mopane

suggests a high water stress in the dry season (February et al.

2007), the species is able to survive water-stress conditions

because of its ability to use water efficiently (Mantlana

2002), which is probably the result of its osmotic adjustment.

By using a combination of physical and _ physiological

adaptations, mainly involving roots and leaves, mopane

is able to tolerate hot, dry conditions mainly found in low-

lying areas of southern Africa.

All these adaptation mechanisms enable mopane to use

the available limited moisture and nutrients efficiently in

order to survive semi-arid to arid conditions of southern

Africa. This article has further reviewed and tested the effect

of environmental factors on the distribution of mopane

in southern Africa. The variables used include rainfall,

temperature, altitude, and soil types.

Environmental drivers and factors

influencing mopane distribution

Various authors have shown that the distribution of mopane

in southern Africa is associated with climatic and edaphic

factors (e.g. Madams 1990; Mapaure 1994; O’Connor 1992;

Werger & Coetzee 1978). Its distribution is principally

influenced by moisture availability expressed through

rainfall, temperature, altitude and soil texture (Bennett 1985;

Henning 1976; Mapaure 1994; Stevens et al. 2014; Timberlake

1995; Werger & Coetzee 1978). It should be noted that rainfall

and temperature co-vary with altitude; however, Stevens et al.

(2014) indicated that there is little evidence of which factors,

or combinations thereof, determine the distribution limit of

this species. This article therefore reviews and discusses the

extent to which rainfall, temperature, altitude and soil types

influence the distribution of mopane in southern Africa.

The data used were derived from various sources (Table 1

and Table 2). The areas covered by mopane in southern

Africa (Table 1) were plotted against minimum, average

and maximum rainfall, temperature and altitude using a

polynomial regression analysis (Figures 2-4). The rationale

for using polynomial regression was that all functions

(linear and non-linear) were showing weak relationships

and this was worse when fitting a linear function. However,

a polynomial function gave a better fit compared to linear

function, which is the reason it was used here.

Rainfall

Mopane is distributed along a variable rainfall gradient,

ranging from an annual average of 50 mm in Namibia to

1000 mm in Zambia (Table 2). However, areas receiving low

to moderate rainfall, especially between 400 mm and 800 mm

per annum, better correlate with the distribution of mopane

in southern Africa (Madams 1990; Mapaure 1994; Werger

& Coetzee 1978). The above finding is closer to the average

of 369 mm — 700 mm per annum as estimated in this article

(Table 2). It includes all countries within its distribution, with

the exclusion of Zambia where rainfall can reach 1000 mm

http://www.abcjournal.org/ 13 | doi:10.4102/abc.v44i1.152

Review BEE) mm

per annum. However, it should be noted that areas receiving

250 mm — 450 mm of rainfall per annum are considered as

the most favourable environmental niche for the growth and

distribution of mopane (Siebert 2012; Thompson 1960).

& MaxR

AverageR

— — Poly. (MinR)

Poly. (MaxR)

—--— Poly. (AverageR) |

120 000 =

€ 100 000 |

a 80 000 +

@ 60000 -

° J 2

o nee aS Neg pees

S J & é a =e

& 20000 BMAD

os gr??

0 . : r 1 t

0 200 400 600 800 1000 1200

Rainfall (mm)

Source: Authors’ own creation

Note: The dotted and solid lines represent the fitted polynomial regression curve at 95%

confidence interval.

R, rainfall; min; minimum; max, maximum; poly, polynomial regression.

FIGURE 2: Effect of rainfall on the distribution of mopane in southern Africa.

= MaxT

& AverageT

— — Poly. (MinT)

Poly. (MaxT)

---— Poly. (AverageT)

140 000

=~ 100000: i SO a

mo} xy !

® 3800001 alas : ‘

oO = eee /

> 60000; R?=0.0054 \ 2 A,

8 A ‘2 = 0.1367 = 0892

« 400004 &

ov &

= 200001 ¢ dy a

¢ & |

0 Tt T T T T T T

(0) 5 10 15 20 25 30 35

Mean daily temperature (°C)

Source: Authors’ own creation

Note: The dotted and solid lines represent the fitted polynomial regression curve at 95%

confidence interval.

T, temperature; min; minimum; max, maximum; poly, polynomial regression.

FIGURE 3: Effect of temperature on the distribution of mopane in southern

Africa.

@ Minalt

& MaxAlt

& AverageAlt

— — Poly. (MinAlt)

Poly. (MaxAlt)

—--—— Poly. (AverageAlt)

120 000

@ & R?=0.281

ie 100 000 + x

= SS

= 80 000 + A

® 60000 ’

fo)

° 40000

= 20000 | ¢ 4 &

0 T T T T T T

(0) 200 400 600 800 1000 1200

Altitude (m.a.s.I.)

Source: Authors’ own creation

Note: The dotted and solid lines represent the fitted polynomial regression curve at 95%

confidence interval.

Alt, altitude; min; minimum; max, maximum; poly, polynomial regression.

FIGURE 4: Effect of altitude on the distribution of mopane in southern Africa.

The review showed that low rainfall positively correlates

with the distribution of mopane in southern Africa, but the

relationship becomes weak when the annual rainfall exceeds

600 mm per annum (Figure 2). This corroborates well with

the findings by Porter (1968) and Henning (1976), who

both indicated that an increase in rainfall to > 800 mm per

annum becomes the limiting factor to mopane distribution.

As also indicated by Thompson (1960), the limitation of

mopane from higher rainfall zones is probably the result

of competition with other species, which are more suited

to those wetter conditions, low temperature, acidic soil

conditions and high frequency of disturbances such as fires.

However, our analysis indicated that the relationship

between rainfall and mopane distribution was higher at

minimum or low annual rainfall (R* = 0.38), slightly declined

at average rainfall (R* = 0.32) and then significantly declined

at maximum or higher rainfall (R° = 0.28). Although this

relationship is positive, it clearly gives a less than 40%

confidence (Figure 2), which concurs with Stevens ef al.

(2014) that rainfall alone cannot be considered as the major

factor determining the distribution of mopane. It is further

indicated that the probability of mopane presence drops to

< 50% when precipitation exceeds 380 mm in the wettest

quarter (Stevens ef al. 2014), which confirms that the species

favours low rainfall areas (Figure 2). The possibilities of

rainfall decline as a result of climate change means that it

will further favour the distribution of mopane in areas such

as Zambia, which is currently considered a high rainfall area.

Temperature

Mopane is distributed in hot and dry environments, where

temperatures can exceed 35 °C (Dye & Walker 1980; Mucina

& Rutherford 2006; Porter 1968). Low winter temperature

and frost are important limiting factors for mopane

distribution, especially along its southernmost boundary

(Cole 1986; Henning 1976; Siebert 2012; Stevens et al. 2014;

Werger & Coetzee 1978; White 1983). The mean daily

temperature regime within its distribution in southern Africa

ranges from 12 °C to 31°C (Table 2), averaging between 16 °C

and 29 °C, as also found by Du Plessis (2001). However, its

distribution is limited in areas where the average minimum

winter temperature is below 5 °C (Cole 1986; Henning 1976;

Rutherford ef al. 2006; Stevens et al. 2014; Voorthuizen 1976;

Werger & Coetzee 1978; White 1983), which confirms that

the species is adapted to high temperature areas.

The review revealed that the distribution of mopane is best

associated with an average daily maximum temperature

of 30 °C (R* = 0.42) (Figure 3), but that relationship

declines when the mean daily maximum temperature

drops (Figure 3). Although the relationship between mean

daily temperature and mopane distribution is positive,

it clearly gives a less than 43.0% confidence (Figure 3).

Stevens ef al. (2014) also found that minimum temperature

in the coldest month was the strongest determinant for

mopane distribution, accounting for 42.2% of the modelled

distribution. However, these results do not give a degree of

http://www.abcjournal.org,

confidence of at least > 50.0% to better explain an important

factor associated with the distribution of mopane. This

therefore means that temperature alone cannot be considered

as the most important factor determining the distribution of

mopane in southern Africa. However, it is important to take

into account that the probability of mopane presence drops

below 50.0% at minimum temperatures less than 5 °C in the

coldest month of July. Therefore, minimum temperature is

predicted to limit the distribution of mopane from entering

the cold interior of the southernmost boundary of southern

Africa (Stevens et al. 2014). The limitation of mopane at low

temperature zones is because of occasional events of frost

(Stevens et al. 2014; Whitecross et al. 2012), which mainly

destroys trees and shrubs less than 4 m in height (Whitecross

et al. 2012). However, an increase in temperature will further

facilitate the distribution of mopane in areas currently

considered as cold, especially in areas west and slightly

south of its current distribution range (Stevens e¢ al. 2014).

Altitude

Sebego (1999) indicated that topographic location could be

one of the important factors determining the distribution

of mopane. Siebert (2012) found that the occurrence of

mopane is associated with low-lying, flat and undulating

areas. The distribution of mopane is normally along the

flood plains and valley bottoms of large rivers such as the

Cunene, Chobe, Limpopo, Luangwa, Okavango, Shire and

Zambezi (Cole 1986; Mapaure 1994; Werger & Coetzee 1978).

However, this article confirms that mopane can be found at

variable altitudes ranging from 100 m.a.s.l. to 1200 m.a.s.l.

(Table 2), but attaining optimal distribution and growth at

altitudes ranging from 313 m.a.s.1. to 856 m.a.s.l. on average

(Table 2), which is closer to the 200 m.a.s.l. — 600 m.a.s.1.

and 400 m.a.s.l. — 700 m.a.s.l. average as indicated by White

(1983) and Mapaure (1994), respectively. Porter (1968) further

indicated that mopane rarely occurs at altitudes > 900 m.a.s.L.,

but this article has shown that it can be occasionally found at

> 900 m.a.s.1. in countries such as Angola, Namibia and Zambia

(Table 2). These are normally areas where unfavourable soil

conditions prevent the growth of other species (Henning

1976), thus favouring the distribution of mopane.

The relationship between altitude and mopane distribution

is positive; although, that relationship gives a < 45%

confidence (Figure 4). This also means that altitude alone

cannot be considered as the important factor determining

the distribution of mopane in southern Africa. However,

the distribution of mopane correlates well at low altitude

(R* = 0.44), but that relationship declines at higher altitudes

(R° = 0.28) (Figure 4). This finding corroborates the

findings from various authors who also indicated that the

distribution of mopane is associated with low-lying, flat and

undulating areas (e.g. Werger & Coetzee 1978; Cole 1986;

Mapaure 1994; Siebert 2012). This implies, as also indicated

by Henning (1976), that limited distribution of mopane at

higher altitudes might be the result of combined influences

of increased precipitation, lower temperatures, acidic soils

and disturbances such as fires.

doi:10.4102/abe.v44il.152 a a wn

Soil type

According to Madams (1990), soil type also correlates well

with the distribution of mopane. Mopane is capable of

surviving on a variety of soil types (Henning 1976; Madams

1990). It grows in arid areas on relatively fertile, fine-grained

soil, sandy-loamy soil to clayey soil (Henning 1976; Madams

1990; Timberlake 1995; Werger & Coetzee 1978) (Table 2).

Mopane is most frequently associated with shallow soils

(Henning 1976) and normally found in alluvium and

colluvium soils (Timberlake 1995). It is sometimes found

in deep soil (Mapaure 1994; Werger & Coetzee 1978), but

the calcrete layer near the surface hinders mopane root

penetration into the deep soil.

The soils in areas where mopane occurs tend to have high

exchangeable sodium content (Dye & Walker 1980; Werger

& Coetzee 1978), which inevitably results in reduced

permeability and increased susceptibility to soil erosion

(Scholes 1997). Mopane mainly survives on alkaline soils

(Werger & Coetzee 1978) and is less common on acidic

soils (Henning 1976). White (1983) further indicated that

mopane does not occur on true saline soils in which water-

soluble salts exceed 0.2% — 0.3%. As a result, mopane is thus

considered as an indicator species of alkaline soil (Werger &

Coetzee 1978).

Soil having low nitrogen (< 0.2% at 0 cm —- 10 cm),

phosphorus (< 1.5 ppm), low moisture (15.0%) and

exchangeable magnesium favours the growth and

performance of mopane, but an increase in soil sodium

and potassium levels results in a decline in the growth

yield, which is probably because of increased soil osmotic

suction, whilst increasing magnesium seems to improve

soil moisture uptake (Henning 1976). Therefore, mopane

exhibits a shrub structure on shallow sodium-rich soils

or clay soils derived from basalt (Mapaure 1994; Mlambo

2006). These are areas with limited soil depth and are

normally occupied by ‘bonsai’ shrubby mopane which

grow up to 1.5 m in height. The ‘cathedral’ mopane grow

quite tall on deep nutrient-rich alluvial soils (Mapaure

1994; Timberlake 1995): up to 6 m in height on heavy

0.6 ®

Mozambique

0.5 4

0.4 5 i Malawi

South Africa i @

N e Cots

cS eee e ©

2 age e@ Zimbabwe

Ss 0.2 4 Namibia

(a)

0.1 5

0.0) i : ae

Botswana

-0.1 + 7 T r ; —

-0.2 0.0 0.2 0.4 0.6 0.8 1.0

DCA Axis 1

Source: Authors’ own creation

DCA, detrended correspondence analysis.

FIGURE 5: Ordination of the drivers of mopane distribution in different

countries in southern Africa.

impervious soils and up to 25 m in areas having sandy-

loamy and alkaline soils (Werger & Coetzee 1978). It is also

important to note that the distribution of mopane is limited

in the acrisolic soils, possibly because acrisols derived

from acid igneous and metamorphic rocks, limit the

growth of mopane, but support the growth of other species

such as Acacia, Boscia, Grewia, Combretum and Terminalia

(Madams 1990).

Ordination of mopane distribution

An ordination diagram illustrates the environmental factors

included in Table 2 that are suggested to influence the

distribution of mopane in different countries in southern

Africa (Figure 5). The ordination graph shows that mopane

is distributed in variable environmental conditions

in different countries, but there are some _ similarities.

For instance, mopane in Namibia occurs under similar

environmental conditions to those in Angola, but quite

different from those in Mozambique, Malawi and Botswana

(Figure 5). In addition, mopane in South Africa occurs under

similar environmental conditions to those in Zimbabwe

and Zambia, but quite different from those in Angola,

Namibia, Mozambique, Malawi and Botswana (Figure 5).

However, mopane demonstrates a relatively wide tolerance

range for the various environmental factors under which it

occurs within its distribution range.

Conclusion

Colophospermum mopane is distributed along variable local

climatic, topographical and edaphic factors in the low-lying

areas of -southern Africa. It mainly occupies areas receiving

low to moderate rainfall, at low lying altitudes, with high

temperature and variable soil types. An increase of annual

rainfall (> 800 mm), altitude (> 800 m.a.s.l.), acidic soil and

a decline in minimum winter temperature (< 5 °C) limits the

distribution of mopane. Limited distribution of mopane in

areas receiving high rainfall, low temperatures and at higher

altitudes is probably a result of the combined effects of freeze

events, competitive interactions with other species and

disturbances such as fires. An increase in temperature has

the potential to drive mopane from its current distribution in

high temperature areas to colder zones at its southernmost

boundaries, whilst a reduction in annual rainfall could drive

mopane from its current distribution in low to moderate

rainfall areas to high rainfall zones. However, this article

demonstrated that the distribution of mopane in southern

Africa is not fundamentally determined by climatic factors,

but possibly by edaphic factors (soil type and nutrients),

competitive interaction with other species and disturbances

such as fires and browsing activity by large herbivores.

It is further concluded that the physical, chemical and

physiological responses of mopane enable it to survive

various disturbances and ‘harsh’ environmental conditions

in southern Africa’s savanna ecosystem. This means that

a better understanding of the adaptation mechanisms

and distribution of mopane is critical and can be used to

= http://www.abcjournal.org/ 15 | doi O41 02/abev44i1152) = == aaa

explain the distribution and survival of the species in these

‘harsh’ conditions in southern Africa. This understanding

can also be used to further identify the ecology of the

many mammalian and invertebrate herbivores that are

found within the mopane ecosystem. Such information is

essential for holistic management of mopane woodland and

shrublands in southern Africa.

However, because of the complexity associated with

identifying factors which associate best with the distribution

of mopane in southern Africa, we recommend that such

complexity be addressed through the development of an

integrated model. Such a model needs to include climatic

factors (e.g. rainfall and temperature), topographical factors

(e.g. altitude and slope), edaphic factors (e.g. soil types and

soil nutrients) and disturbances (e.g. fires, herbivory and

competition). Once developed, such a model can significantly

improve the precision of predicting the distribution of not

just mopane, but also other vegetation formations and

associated wild animals in the savannas.

Acknowledgements

We would like to acknowledge the comments and

suggestions provided by Prof. Laco Mucina from the

University of Western Australia and Prof. Jonathan R.

Timberlake from the Royal Botanic Gardens during the initial

draft of this article. We also appreciate the comments, inputs

and suggestions provided by two anonymous reviewers.

Your contributions assisted a great deal in improving the

quality of this article.

Competing interests

The authors declare that they have no financial or personal

relationships that may have inappropriately influenced

them in writing this article.

Authors’ contributions

R.A.M. (University of Limpopo) was responsible for

reviewing the literature and writing of the article. I.M.

(University of Namibia), M.J.P. (University of Limpopo),

W.J.L-P. (University of Limpopo) and A.T.S. (National

Research Foundation) were responsible for providing

technical input in the article and supervision.

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http://www.abcjournal.org/ doi:10.4102/abe.v44i1.152 — =

= Pp a ge {| oO f 4 S h or t N oO t e reer reer errr etree ee eT eee RTO OS ROLE CLR OTT PEPE PEPE PTR OO TEETER PTET

Moraea orthrosantha (\Iridaceae: Irideae), a new

species from Namaqualand, South Africa

Authors:

Peter Goldblatt!”

John C. Manning??

Affiliations:

1Missouri Botanical Garden,

St. Louis, United States of

America

*Research Centre for Plant

Growth and Development,

University of KwaZulu-Natal,

South Africa

3Compton Herbarium, South

African National Biodiversity

Institute, South Africa

Correspondence to:

John Manning

Email:

j.manning@sanbi.org.za

Postal address:

Private Bag X7, Claremont

7735, South Africa

Dates:

Received: 05 Nov. 2013

Accepted: 25 Mar. 2014

Published: 04 July 2014

How to cite this article:

Goldblatt, P. & Manning,

J.C., ‘Moraea orthrosantha

(Iridaceae: Irideae), a new

species from Namaqualand,

South Africa’, Bothalia 44(1),

Art. #132, 4 pages. http://

dx.doi.org/10.4102/abc.

v44i1.132

Licensee: AOSIS

OpenJournals. This work

is licensed under the

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Attribution License.

Read online:

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Background: Recent fieldwork in Namaqualand, Northern Cape Province, South Africa,

indicated the existence of an undescribed member of Moraea subgenus (subg.) Umbellatae

Goldblatt & J.C.Manning, a small, early branching clade of the genus.

Objectives: To describe the new species of Moraea subg. Umbellatae.

Method: Recent collections were compared with existing material and published literature.

Results: Moraea orthrosantha is described as a new species, differing from Moraea margaretae in

the well-developed nodes between the cauline leaves. Two earlier collections of the species were

misidentified as M. margaretae.

Conclusion: The new species increases our understanding of the diversity of Moraea in southern

Africa and assists in conservation assessments of both M. margaretae and M. orthrosantha.

Introduction

Fieldwork in Namaqualand, Northern Cape Province, South Africa, in late spring of 2012

and 2013 yielded the discovery of a population of a yellow-flowered Moraea, representing an

undescribed member of subgenus (subg.) Umbellatae Goldblatt & J.C.Manning, an early branching

clade of the genus currently comprising nine species (Goldblatt & Manning 2013; Goldblatt,

Manning & Schnitzler 2013). The new species, described here as Moraea orthrosantha, has a well-

developed and multi-branched aerial stem with a solitary leaf at each node, conventional Moraea-

type flowers with larger outer tepals and well-developed, petaloid style branches with prominent

crests. These features are plesiomorphic in the genus (Goldblatt et al. 2013), suggesting that

M. orthorosantha is unspecialised in subg. Umbellatae. The largely sub-Saharan genus Moraea now

comprises some 226 species (Goldblatt & Manning 2013), most of them restricted to the southern

African winter rainfall region. Subgenus Umbellatae now includes 10 species, all endemic to

western South Africa and extending from the Richtersveld of Northern Cape Province to the

Cape Peninsula and Caledon District in Western Cape Province. We include a key to the species

of subg. Umbellatae.

Research method and design

We examined all relevant collections at Bolus Herbarium, University of Cape Town (BOL);

Compton Herbarium, South African National Biodiversity Institute, Cape Town (NBG), National

Herbarium, South African National Biodiversity Institute, Pretoria (PRE) and South African

Museum Herbarium, South African National Biodivoersity Institute, Cape Town (SAM), the

primary southern African herbaria (acronyms after Holmgren, Holmgren & Barnett 1990). Plants

were examined in the field for 3 h and 12 specimens were pressed as type material for distribution

to herbaria (as cited below). Type material was collected under permit number 010/2012 from the

Northern Cape Department of Environment and Nature Conservation. Additional specimens are

cited following the Degree Reference System (Leistner & Morris 1976).

Taxonomic treatment

Moraea orthrosantha Goldblatt & J.C.Manning, sp. nov.

Type: SOUTH AFRICA. Northern Cape: 3017 (Hondeklipbaai):15.5 km N of Garies, sandy slopes

at + 645 m, 30°27’ S 17°37’ E (-BD), 10 Oct. 2013, Goldblatt & Porter 13990 (NBG, holo.; K, MO, PRE,

isO.).

Description

Plants up to 350 mm high. Corm: 8 mm — 11 mm diameter; tunics brown, + woody, inner

layers entire, outer splitting from base. Stem: with three or four (five) internodes, with one or

http://www.abcjournal.org/ doi:10.4102/abe.v44il.132

Page 2 of 4

Short Note

be IE eg

Source: Artist — John Manning

Scale bar: (a) 10 mm; (b) 2 mm.

FIGURE 1: Diagrammatic representation of Moraea orthrosantha holotype depicting, (a) the flowering plant and (b) stamens and style.

http://www.abcjournal.org/ @tReE doi:10.4102/abc.v44i1.132

= a cr erernctmncnmermmrmrreress EDDY ge 3 0f 4 a SHO NOC mr nn nnn ere eee

two branches at each node; branches slightly flexed above

sheath of subtending leaf. Leaves: three to five, lowermost

longest, inserted 10 mm — 20 mm above ground, linear,

channeled, + straight or loosely twisted, up to 450 mm long,

upper leaves progressively shorter, none entirely sheathing.

Rhipidial spathes: inner 35 mm — 40 mm long, + truncate,

apex brown, somewhat lacerated; outer half to two-thirds as

long, with acute, brown apex. Flowers: buff-yellow, lightly

brown-veined, outer tepal limbs with deep yellow nectar

guides at bases edged brown, darkly veined abaxially,

spreading slightly below horizontal, inner tepals with

reddish-brown spot at bases of limbs, similarly spreading;

outer tepals oblanceolate, 30 mm — 32 mm long, limbs

20 mm — 22 mm x 10 mm — 12 mm, claws + 10 mm long;

inner tepals oblanceolate, + 22 mm x 5 mm — 6 mm.

Stamens: with filaments 8 mm — 9 mm long, united in lower

5 mm — 6 mm, diverging distally; anthers dark purple,

3.5 mm — 5.0 mm long; pollen red. Ovary: exserted, oblong-

elliptic, 5 mm — 6 mm long; style branches + 10 mm long,

crests 10 mm — 14 mm long, erect, narrowly wedge-shaped.

Capsules: narrowly obovoid, 9 mm — 11 mm long. Seeds: softly

angular, + 1 mm long, with rugulo-reticulate surface, light

yellow-brown. Flowering time: late September to late October;

flowers opening + 07:00 and collapsing + 12:00 (Figure 1).

Distribution and ecology

Moraea orthrosantha is known only from a small area north

of Garies in Northern Cape Province, South Africa, in the

higher country south of Garagams (Figure 2). Plants appear

to be restricted to locally wetter sites; in trampled or drier

situations they are much smaller in size. The soil in which

they grow is the typical granitic gravel of Namaqualand and

plants sometimes occur in rocky ground. We did not make

an accurate count of the number of plants at the site but we

BOX 1: Key to the species of Moraea subgenus Umbellatae.

1A. Plants with stem subterranean; leaves and rhipidia clustered at ground level:

2A. Tepals unequal, outer larger and with nectar guides at limb bases; style branches well developed with prominent crests...

2B. Tepals subequal, outer + as long as inner and both with nectar guides at limb bases; style branches divided almost to base into paired

filifonmianmsiwithoUticrestsis) Myce ea EL ANN A ated sioaia but a utamen mips Als Os eile da it By a ean DRv INURE acne SO ALR a On es es CLS Rau ee

1B. Plants with stem aerial; either lowermost leaf inserted at ground level or all leaves and branches clustered at ground level but then rhipidia stalked:

3A. Foliage leaves all clustered at ground level; branches (when present) produced at ground level f ne . M. margaretae

3B. Plants wither with lowermost leaf inserted at or close to ground level or all leaves inserted at stem apex well above ground; branches

(when present) arising well above ground:

4A. Lowermost leaf (of two or more) inserted at or close to ground level:

SA. Branches crowded at stem apex; tepals free

5B. Branches borne along stem, not crowded above:

6A. Tepals free to base; inner tepals present.......

6B. Tepals united in short tube; inner tepals lacking. M. cooperi

4B. All leaves clustered at stem apex well above ground:

7A. Style branches filiform and extended between stamens, WithOUt..........0 ncaa Soe eer ; recon one M. nana

7B. Style branches flattened, appressed to opposed anthers, with or without prominent crests:

8A. Style branches bearing prominent crests 10 mm — 12 mm long; filaments united in

column 8 mm — 10 mm long

8B. Style branches without crests; filaments in column 5 mm—7 mm long, united or free distally:

9A. Plants 150 mm — 450 mm high; outer tepals 19 mm — 27 mm x 9 mm — 12 mm; inner rhipidial

spathes 30 mm —40 mm long.

9B. Plants 90 mm — 150 mm high; outer tepals 15 mm — 24 mm x 6 mm— 10 mm; inner rhipidial spathes

20 mm — 35 mm long.

located over 40 individuals, including some immature and

not of flowering age.

Etymology

As the specific epithet suggests, flowers open soon after

sunrise and collapse at about noon. The epithet derives from

the Greek orthros [morning] and anthos [flower]. There are

few other examples of this flowering phenology in the genus

and none known in subg. Umbellatae.

Conservation status

The habitat is relatively undisturbed, although it is located

less than 20 m from the main road from Cape Town to

Springbok. Future road expansion will severely impact the

(©) Moraea orthrosantha

©) Moraea margaretae

16° 18° 20° 2pe? 24°

FIGURE 2: Distribution of Moraea orthrosantha and the closely related Moraea

margaretae.

M. longiflora

M. singularis

...M. linderi

eek M. orthrosantha

..M. intermedia

...M. umbellata

..M. maximiliani

http://www.abcjournal.org/ 20 | toy Moy /itopy fella Valu sy)

known habitat. Summer grazing by stock should not affect

the population as the plants will be dormant. The possible

occurrence of additional populations in the area requires

further investigation.

Diagnosis

In its vegetative and floral morphologically, M. orthrosantha

is perhaps the least specialised member of subg. Umbellatae.

The species is distinguished by its vegetative habit: mature

plants have stems up to 350 mm high with several leaves,

one at each of up to four or five aerial nodes and one or

two branches per node. The flowers are typical of the

genus, having free tepals, well-developed petaloid style

branches with erect crests and prominent nectar guides at

the bases of the outer tepal limbs. Somewhat unusually,

the bases of the inner tepal limbs are marked with a small,

dark reddish-brown spot. The buff or dull yellow colour

of the perianth is typical of the subgenus. In contrast, most

other species of the subgenus have branches crowded at the

first aerial node, this either approximately at ground level

(Moraea margaretae Goldblatt) or well above ground level

(Moraea linderi Goldblatt and Moraea intermedia Goldblatt

& J.C.Manning). All three of these species have well-

developed style branches and crests but other species have

the style branches reduced to narrow lobes without crests

[Moraea maximiliani (Schltr.) Goldblatt & J.C.Manning and

Moraea umbellata Thunb.] or divided into paired filiform

arms [Moraea nana (L.Bolus) Goldblatt and Moraea singularis

Goldblatt & J.C.Manning]. The remaining two species

(Moraea cooperi Baker and Moraea longiflora Ker Gawl.) have

the tepals united in a perianth tube.

The morning blooming phenology is unusual in Moraea:

the flowers of most species of subg. Umbellatae for which

phenology is known open in the late morning and collapse

in the late afternoon (exact times are unrecorded for any

of these species). Records for M. margaretae, with which

M. orthrosantha is most easily confused, indicate that

the flowers open mid-morning and collapse toward

sunset (Goldblatt 1986). Perhaps significantly, the three

Namaqualand species of Moraea sect. Flexuosae Goldblatt,

including Moraea schlechteri (L.Bolus) Goldblatt, also have

fugaceous flowers opening early in the morning and

collapsing after 12:30.

SHOP NOC mr rere trent woes oee

History

Moraea orthrosantha was first recorded by Rudolf Schlechter

in 1897 at Brakdam between Garies and Garagams (south of

Kamieskroon) in Namaqualand. His collection and a subsequent

one made by Frances Leighton in 1945 constitute the only early

records of the species. Both were referred to as M. margaretae

(Goldblatt 1976), an action clearly mistaken in light of the

new collections and examination of living plants. These early

collections bracket the type locality, 15.5 km north of Garies.

Additional specimens examined

SOUTH AFRICA. Northern Cape: 3017 (Hondeklipbaai): hills

at ‘Brackdam’, (—BD), 08 Sept. 1897, Schlechter 11120 (BM);

6 miles [+ 9 km] north of Garies (-DB), 03 Sept. 1945, Leighton

1129 (BOL).

Acknowledgements

We thank Lendon Porter for his assistance and companionship

in the field and Northern Cape Department of Environment

and Nature Conservation for the collecting permit to J.C.

Manning.

Competing interests

The authors declare that they have no financial or personal

relationships that may have inappropriately influenced them

in writing this article.

Authors’ contributions

Both authors collaborated on all aspects of the research.

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from the Northern Cape, South Africa

Authors:

Nicola G. Bergh?

Nick A. Helme?

Affiliations:

1Compton Herbarium, South

African National Biodiversity

Institute, South Africa

2Nick Helme Botanical

Surveys, Scarborough,

South Africa

Correspondence to:

Nicola Bergh

Email:

n.bergh@sanbi.org.za

Postal address:

Private Bag X7, Claremont

7735, South Africa

Dates

Received: 20 Feb. 2014

Accepted: 04 July 2014

Published: 18 Sept. 2014

How to cite this article:

Bergh, N.G. & Helme, N.A.,

2014, ‘A new species of

Berkheya (Asteraceae,

Arctotideae) from the

Northern Cape, South

Africa’, Bothalia 44(1), Art.

#123, 5 pages. http://dx.doi.

org/10.4102/abc.v44i1.123

Licensee: AOSIS

OpenJournals. This work

is licensed under the

Creative Commons

Attribution License.

Read online:

ure Scan this OR

Els #0] code with your

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Eline * toread online.

Background: Berkheya is a large, mainly southern African genus of approximately 75 species,

several of which are poorly known and under-collected. Since revision in 1959, only a few new

species have been described. Publication of new taxa facilitates conservation objectives and

contributes to a better understanding of the southern African flora.

Objectives: The objectives of this study were to describe a new species of Berkheya, consider

its taxonomic position within the genus and assess its conservation status.

Methods: Macromorphology and micromorphology of the new species were compared with

known species.

Results: Berkheya dumicola N.G.Bergh & Helme was described from two subpopulations from

the northern Bokkeveld escarpment, Northern Cape Province, South Africa. The species is a

tall shrub with radiate flowerheads, toothed receptacle alveole margins, a uniseriate pappus

of small, rounded scales and very short twin-hairs on the cypsela.

Conclusion: Berkheya dumicola is a new species with a unique combination of features. Based

on morphological characteristics, its closest relative within the genus is likely to be the recently

described Berkheya chrysanthemoides J.C.Manning & Goldblatt. The limited geographic extent

and small population size of B. dumicola warrant an International Union for Conservation of

Nature (IUCN) status of ‘Endangered’.

Introduction

Berkheya Ehrh. (Asteraceae) is the largest genus in Arctotideae subtribe Gorteriinae, the ‘spiny

daisies’ of southern Africa. The genus has been shown to be paraphyletic using both molecular

(Funk & Chan 2008; Funk, Chan & Keeley 2004) and morphological data (Karis 2006) and changes

in generic circumscriptions in the subtribe are anticipated (Karis et al. 2009). Based on current

evidence, Berkheya consists of approximately 75 species distributed predominantly in South

Africa. The genus was last revised by Roessler (1959) and has thus not been treated as a whole

for over 50 years, despite several useful regional treatments, including Goldblatt and Manning

(2000), Hilliard (1977), Manning and Goldblatt (2012a) and Snijman (2013), and the description

of a handful of new species (Hilliard 1977; Hilliard & Burtt 1975; Manning et al. 2010; Manning

& Goldblatt 2012b). The prevalent covering of sharp spines makes Berkheya species difficult

and painful to collect and the genus is poorly known and under-collected, despite forming a

prominent component of many plant communities.

Berkheya species occur in several bioregions but are concentrated in the Greater Cape Floristic

Region, where they are most strongly associated with the winter-rainfall desert (succulent Karoo)

vegetation (Verboom ef al. in press). Most species are perennial herbs, less commonly shrubs,

distinguished from allied genera by the possession of, (1) involucral bracts that are always spiny

and basally connate but never hardened into a woody structure, (2) a honeycombed receptacle

with the cypselas embedded in alveoli that are all of equal size and (3) a pappus (Roessler 1959).

Roessler (1959) divided Berkheya into eight series based on characters of the receptacle alveolar

margins, cypsela hair type, pappus arrangement and structure, degree of connation of

the involucral bracts, leaf morphology (including degree of division, margin serration and

indumentum), leaf arrangement (opposite versus alternate), presence or absence of ray florets

and growth form. Additional characters used to define species include hairiness of the stems

and leaves, leaf shape and size, form of the leaf margins, size and structure of the leaf spines,

arrangement and size of the capitula, ray floret colour and the size and morphology of the

involucral bracts.

Berkheya material from a localised area in the northern Bokkeveld escarpment in the Calvinia

region of the Northern Cape Province (Figure 1) was examined and this could be distinguished

from other species in the genus by its tall, shrubby habit, relatively large and broad leaves, small

doi:10.4102/abce.v44i1.123

radiate heads clustered in paniculate groups, very short

uniseriate pappus scales and short hairs on the cypselas.

This material represents a new species which is described

in detail here.

Research method and design

Berkheya specimens from the South African National

Biodiversity Institute’s (SANBI) Compton Herbarium in Cape

Town (NBG), SANBI’s South African Museum Herbarium in

Cape Town (SAM) and the Bolus Herbarium at the University

of Cape Town (BOL) were compared with those of the new

taxon. Holotype material was collected and dried using

standard techniques in October 2013. Type material has been

deposited in NBG, BOL and SANBI’s National Herbarium

in Pretoria (PRE), Kew Herbarium at the Kew Royal Botanic

Gardens, United Kingdom (K) and the Swedish Museum of

Natural History Herbarium in Stockholm (S).

Habitat and habit were documented in the field.

Morphological structures were examined on dried and

rehydrated specimens using a dissecting microscope. Images

were captured from the microscopes using an Olympus SC30

digital camera attachment and the software Analysis getIT

v. 5.1 (Olympus Soft Imaging Systems, Miinster, Germany).

Measurements were made on digital images using MeasurelT

v. 5.1 from the same company.

Plants were collected under collecting permit FLORA

02/02/2013 issued to Compton Herbarium staff by the

Department of Nature Conservation, Northern Cape Province.

Taxonomic treatment

Berkheya dumicola N.G. Bergh & N.A. Helme, sp. nov.

Type

SOUTH AFRICA. Northern Cape: 3119 (Calvinia): Noord

Bokkeveld, top of Die Hel pass, north-west of farm

Height above

32° sea level (m)

[___] 300

Source: Map drawn by M. Smith

The black dot represents the location of the two known populations of Berkheya dumicola.

FIGURE 1: Distribution of Berkheya dumicola.

http://www.abcjournal.org/ 23 | doi:10.4102/abc.v44i1.123

- Page 2 0f 5 eo ST EN

Kookfontein (-AA), 12 Oct. 2013, Helme 7794 (NBG, holo.;

BOL, PRE, K, S, iso.).

Description

WelLbranched shrub up to 2 m tall, with gnarled woody

stem to 120 mm diameter at base, branches leafy towards

apices (Figure 2a), thinly (glandular-) tomentose. Leaves:

alternate, sessile, oblanceolate in outline, 30 mm — 80 mm x

20 mm — 50 mm, lamina narrowed to slender base + 2 mm

wide, pinnatifid, two-jugate with five primary lacinia,

lacinia increasing in size distally, primary lateral lacinia

each with smaller secondary lobe in distal axil, narrowed

at base and often with additional slender patent, lobes

resembling spines, lacinia somewhat concave, triangular

to narrowly triangular, shorter than to as long as the

width of undivided portion, excurrent in apical yellowish

spine but lacking spines along margins, margins revolute

(Figure 2b), discolorous, adaxial surface thinly woolly when

young, glabrescent, abaxial surface densely white-felted.

Capitula: shortly pedunculate, 3-10 in loose corymbs at

ends of short, slender flowering branches, several branches

arranged together in paniculate synflorescences, radiate,

20 mm — 35 mm diameter across expanded ray florets,

florets rich yellow. Involucral bracts: 4-seriate (Figure 2c),

basally connate ininvolucre+4mm deep, squarrose, concave,

narrowly lanceolate, margins thickened, cartilaginous,

yellowish, with apical spine 2 mm — 3 mm long and with

2 or 3 pairs of similar marginal spines, glabrous, outer

bracts 5 mm — 8 mm x 2 mm — 3 mm, median 10 mm —

12 mm x 2 mm — 3 mm, inner narrowly lanceolate, 6 mm —

8 mm x 1.5 mm — 2.0 mm. Receptacle: deeply alveolate

(Figure 2d), ovaries embedded in fleshy alveolar tissue,

alveole margins extended into irregularly serrated

cartilaginous teeth 1.0 mm —- 2.5 mm long (Figure 3d).

Ray florets: 8-10, sterile and lacking pappus, attached to

small alveoles fused to inner surface of innermost involucral

bracts (Figure 2d), tube + 8 mm long, glandular-pubescent,

limb narrowly oblanceolate, 12 mm-15mm x 3mm-5mm,

lamina apex unevenly 4-toothed with central division much

shallower than two lateral divisions (Figure 3a). Disc florets:

16-18, hermaphrodite, corolla funnel-shaped, 8 mm -9 mm

long, tube + 5 mm long, densely glandular-pubescent with

long pale hairs, deeply lobed, lobeserect, narrowly lanceolate,

+ 4 mm long, apically sparsely hairy with short clavate

orange hairs on abaxial surfaces, lobe margins somewhat

thickened (Figure 3b). Anthers: sagittate with lanceolate

apical appendage, + 5 mm long (Figure 3c). Style: papillose,

with indistinct ring of antrorse hairs well below the

branching point, style branches tapering towards apex,

obtuse (Figure 3c). Cypselas: obovoid, 2.0 mm — 3.0 mm x

0.8 mm, angled, distally densely covered with, short,

antrorse twin-hairs (Figure 3e). Pappus: of short scales,

uniseriate, + 20, entire, + 0.3 mm x 0.2 mm, oblong-ovate,

apex obtuse-truncate, glabrous (Figures 3e and 3f). Pollen:

lophate. Flowering time: September to October.

Distribution and habitat

This large, shrubby species has been collected from only

one locality in the northern Bokkeveld, north-west of

Nieuwoudtville, on the very edge of the escarpment

(Figure 1). Two subpopulations, each of approximately

30 plants, have been found growing about 800 m apart on

steep, southwest-facing slopes amongst large sandstone

boulders (Figure 2a). The plants occasionally sprawl over the

rocky outcrops. The aspect appears to be important because

B. dumicola was not seen in otherwise similar habitats along

the escarpment edge which generally have a drier, more

northerly aspect.

Ecology

The species grows in tall, fire-protected thicket vegetation.

It is estimated that the habitat has not burnt for at least

80 years. The plants are rooted in deep loamy soils

and, because of the steep slope and dense surrounding

vegetation, are often partly shaded for part of the day. The

thicket vegetation in which they occur includes species

such as Kiggelaria africana L., Gymnosporia buxifolia (L.)

Szyszyl., Lobostemon glaucophyllus (Jacq.) H.Buek., Stachys

sp., Podalyria myrtifolia (Retz.) Willd. and Diospyros austro-

africana De Winter. Average annual rainfall in the area is

= Page 3 0f 5 Short Notes

Source: Photographs (a) and (b) taken by N. Helme, (c) and (d) taken by N. Bergh, of

rehydrated material from the type collection

FIGURE 2: Macromorphological features and habitat of Berkheya dumicola,

including, (a) habitat and habit, (b) inflorescence comprising clusters of three

to ten heads; large leaves with spines present at the lacinia apices but not along

the margins, (c) base of capitulum showing short peduncle and spiny, 4-seriate

involucre and (d) deeply alveolate receptacle with irregularly fimbriate alveolar

margins (disc florets corollas removed).

likely to be in the order of 400 mm — 500 mm per year, with

pronounced and rapid declines to the east and west of the

escarpment edge (Manning & Goldblatt 1997).

Etymology

The specific epithet ‘dumicola’ means ‘thicket dweller’

(Stearn 1967) and refers to the rocky, fire-protected vegetation

in which this species occurs.

Diagnosis and relationships

Berkheya dumicola is an unusually large, shrubby species in

a genus dominated by perennial herbs. The shrubby habit,

radiate heads and relatively broad leaves (Figure 2b) are

most consistent with Berkheya series Fruticosae Roessler,

but members of this series are characterised by entire

or shortly toothed (rarely fringed) receptacle alveolar

margins, densely hairy cypselae with the silky hairs often

long (up to 4 mm), and a biseriate pappus of lanceolate

or subulate scales. The conspicuously fimbriate, serrate-

toothed alveolar margins, shortly pubescent cypselae

with twin-hairs and uniseriate pappus of short, obtuse

scales (Figures 3d—3f) set B. dumicola apart from members

of series Fruticosae. Only one other Berkheya species

500 pm

Source: Photographs taken by N. Bergh

FIGURE 3: Micromorphological features of Berkheya dumicola, taken from

rehydrated material of the type collection, depicting, (a) ray floret and

associated receptacular tissue, (b) disc floret, (c) dissected disc floret showing

style and anthers, (d) section through receptacle to show embedded cypselae

and alveolar margins, (e) dry cypsela and (f) pappus.

~~ ~~ — ———— http://www.abcjournal.org/ 24 | doi:10.4102/abc.v44i1.123 =

shares this unusual combination of characters: the

recently described Berkheya chrysanthemoides from the

nearby central Bokkeveld. Berkheya dumicola differs from

B. chrysanthemoides in having leaves that are broader (20 mm —

50 mm in B. dumicola; 15 mm —30 mm in B. chrysanthemoides),

lack marginal spines (both species have spines on the

lacinia apices but B. chrysanthemoides also possesses smaller

antrorse spines along the margins) and are conspicuously

tomentose on their abaxial surfaces (contrasting with the

abaxially glabrous leaves of B. chrysanthemoides). The capitula

of B. dumicola are smaller (20 mm — 35 mm in B. dumicola;

40 mm — 50 mm in B. chrysanthemoides) and clustered together

in larger groups (three to ten in B. dumicola; one to three in

B. chrysanthemoides).

Manning et al. (2010) speculated that B. chrysanthemoides

may have its closest taxonomic relatives in Roessler’s

(1959) series Rigidae, the species of which are characterised

by similar alveole, pappus and cypsela characters, as well

as sharing anther and pollen features. Species in series

Rigidae, however, are generally herbaceous or suffruticose

and lack ray florets. Berkheya chrysanthemoides, and now

B. dumicola, therefore represent unusually large and radiate

putative members of this series. It is remarkable that these

likely sister-species, both only recently discovered, occur

relatively nearby on the Bokkeveld escarpment. Both species

are known from only a single locality each and further

collections are required to determine their full geographical

ranges and morphological variation.

Conservation status

The total global population of B. dumicola is currently

thought to be less than 100 mature plants and the species

thus fits the criteria for EN D1 (International Union for

Conservation of Nature [[UCN] 2001). No threats to the

species are currently known.

Other material examined

SOUTH AFRICA. Northern Cape: 3119 (Calvinia): Noord

Bokkeveld, top of Die Hel pass, north-west of farm

Kookfontein (-AA), 16 Sept. 2006, Helme 4225 (NBG).

Discussion

Berkheya dumicola is morphologically most similar to

B. chrysanthemoides and together they form an unusual

and geographically proximate subgroup within Berkheya

(B. chrysanthemoides is also from the Bokkeveld plateau,

growing in the Oorlogskloof Nature Reserve approximately

50 km to the south of where B. dumicola occurs). Of

consideration is the fact that both species are known from

only very few specimens, so the documented range of

within-species variation is small. However, many species

do occur naturally in very geographically restricted ranges

and with only a small number of individuals. Assessment of

natural variation in these cases must, of necessity, be based

Short NOt@$ ee

on a small number of specimens. Careful comparison of

morphological differences between B. chrysanthemoides and

B. dumicola revealed a suite of characters, derived from both

reproductive and vegetative structures, differing sufficiently

and consistently to warrant separating the two taxa.

Conclusion

Berkheya dumicola is a new, possibly endangered species

recently discovered in the northern part of the Bokkeveld

plateau in the Northern Cape Province of South Africa.

Acknowledgements

Thanks to G.A. Verboom and S. Smuts for fieldtrip assistance

and R.J. McKenzie for a translation of Roessler’s (1959) Latin

key to Berkheya. We are grateful to M. Smith for production

of the map and plates. We also thank S. Magoswana for

assistance with microscopy and J. Manning for helpful

comments on the manuscript. Northern Cape Nature

Conservation provided permits.

Competing interests

The authors declare that they have no financial or personal

relationships that may have inappropriately influenced them

in writing this article.

Authors’ contributions

N.A.H. (Nick Helme Botanical Surveys) discovered the plant

and recognised it as a new species, as well as contributing

to the manuscript and writing the sections on ecology and

conservation status. N.G.B. (SANBI Kirstenbosch) performed

the dissections and microscope work and wrote the initial

taxonomy section and draft manuscript.

References

Funk, V. & Chan, R., 2008, ‘Phylogeny of the spiny African daisies (Compositae, tribe

Arctotideae, subtribe Gorteriinae) based on trnL-F, ndhF, and ITS sequence data’,

Molecular Phylogenetics and Evolution 48, 47-60. http://dx.doi.org/10.1016/j.

ympev.2008.03.035

Funk, V.A., Chan, R. & Keeley, S., 2004, ‘Insights into the evolution of the tribe

Arctoteae (Compositae: subfamily Cichorioideae s.s.) using trnl-F ndhF, and ITS’,

Taxon 53(3), 637-655. http://dx.doi.org/10.2307/4135440

Goldblatt, P. & Manning, J., 2000, ‘Cape Plants: A conspectus of the Cape flora of

South Africa’, Strelitzia 9, South African National Biodiversity Institute, Pretoria.

Hilliard, O.M., 1977, Compositae in Natal, University of Natal Press, Pietermaritzburg.

Hilliard, O.M. & Burtt, B.L., 1975, ‘Notes on some plants of southern Africa chiefly

from Natal: IV: 161: Berkheya pannosa Hilliard’, Notes from the Royal Botanic

Garden Edinburgh 34(1), 77-78.

International Union for Conservation of Nature, 2001, /nternational Union for the

Conservation of Nature red list categories and criteria version 3.1, \UCN Species

Survival Commission, Gland, Switzerland.

Karis, P.O., 2006, ‘Morphological data indicates two major clades of the subtribe

Gorteriinae (Asteraceae-Arctotideae)’, Cladistics 22, 199-221. http://dx.doi.

org/10.1111/j.1096-0031.2006.00109.x

Karis, P.O., Funk, V.A., McKenzie, R.J., Barker, N.P. & Chan, R., 2009, ‘Arctotideae’, in

V.A. Funk, A. Susanna, T.F. Stuessy & R.J. Bayer (eds.), Systematics, evolution and

biogeography of Compositae, pp. 386-410, International Association for Plant

Taxonomy, Vienna.

Manning, J.C. & Goldblatt, P., 1997, Nieuwoudtville: Bokkeveld plateau and Hantam

South African wild flower guide, Botanical Society of South Africa, Cape Town.

Manning, J.C. & Goldblatt, P., 2012a, ‘Plants of the Greater Cape Floristic Region 1:

The core Cape flora’, Strelitzia 29, South African National Biodiversity Institute,

Pretoria.

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——- PageSof5

Manning, J.C. & Goldblatt, P., 2012b, ‘Berkheya jardineana (Arctotideae —

Gorteriinae), a new dwarf perennial from Swartruggens, Western Cape’,

Bothalia 42(1), 57-59.

Manning, J.C., Karis, P.O., Goldblatt, P. & Helme, N.A., 2010, ‘Berkheya

chrysanthemoides and Heterorhachis hystrix, two new species of Arctotideae-

Gorteriinae from the southwestern Cape’, Bothalia 40(2), 185-190.

Roessler, H., 1959, ‘Revision der Arctotideae-Gorteriinae (Compositae)’, Mitteilungen

der Botanischen Staatssammlung Munchen It|, 71-500.

Short Notes ——— SSS ac Sa ea

Snijman, D.A., 2013, ‘Plants of the Greater Cape Floristic Region 2: The extra Cape

flora’, Strelitzia 30, South African National Biodiversity Institute, Pretoria.

Stearn, W.T., 1967, Botanical Latin: History, grammar, syntax, terminology and

vocabulary, Thomas Nelson & Sons, London.

Verboom, G.A., Linder, H.P., Forest, F., Hoffmann, V., Bergh, N.G. & Cowling, R.M., in

press, ‘Cenozoic assembly of the Greater Cape flora’, in N. Allsopp, J. Colville & G.A.

Verboom (eds.), Fynbos: Ecology, evolution and conservation of a megadiverse

region, Oxford University Press, Oxford.

http://www.abcjournal.org/ 26 | dorLOM102/anena 42S

Short Note

Taxonomic status of Pelargonium reniforme Curt.

Authors:

Janine E. Victor"

Mmamphe Aphane?

Affiliations:

1Biosystematics Research

and Biodiversity Collections

Division, South African

National Biodiversity

Institute, South Africa

Correspondence to:

Janine Victor

Email:

j-victor@sanbi.org.za

Postal address:

Private Bag X101, Pretoria

0184, South Africa

Dates:

Received: 24 June 2013

Accepted: 13 June 2014

Published: 10 Oct. 2014

How to cite this article:

Victor, J.-E. & Aphane, M.,

2014, ‘Taxonomic status

of Pelargonium reniforme

Curt’, Bothalia 44(1), Art.

#173, 3 pages. http://dx.doi.

org/10.4102/abc.v44i1.173

Licensee: AOSIS

OpenJournals. This work

is licensed under the

Creative Commons

Attribution License.

Read online:

oF ofa] Scan this QR

ne code with your

=~ 6 © smart phone or

mobile device

et 79) to read online.

Background: Pelargonium reniforme Curt. is a morphologically variable species that many

authors have attempted to split or combine. Confusion relating to the differences between the

two subspecies currently included under P. reniforme has impeded attempts to assess their

conservation status. Pelargonium reniforme is closely related to Pelargonium sidoides; the two

species are indistinguishable when not flowering and their distributions overlap in some areas.

Objectives: With this study, we aimed to clarify the taxonomic status of the two subspecies of

P. reniforme, which has relevance in terms of their conservation status.

Method: Leaf shape, petiole length, internode length and flower colour were assessed

by studying herbarium specimens of the two subspecies of P. reniforme and specimens of

P. sidoides. Living specimens of the two subspecies were also examined in their natural habitat.

Results: The current investigation showed that the morphological characters used to

distinguish the two subspecies of P. reniforme are too variable to separate them. Variation in

some morphological characters may be related to environmental conditions.

Conclusion: The recognition of the two subspecies of P. reniforme as distinct taxa is no

longer justified.

Introduction

There are over 200 species of Pelargonium L’Her. ex Aiton in South Africa and comparatively

few species elsewhere in the world (Vorster 2000). The genus Pelargonium is characterised by

a zygomorphic flower with a nectariferous spur fused to the pedicel, five petals (rarely four or

two), with the posterior petals larger than the anterior petals and 10 stamens, of which only two

to seven are fertile (Van der Walt 1985).

There are 16 sections recognised in the genus (Bakker et al. 2004). Pelargonium reniforme Curt.

is a member of section Reniformia (R.Knuth) Dreyer ex J.P.Roux (Roux 2013) and is distributed

from Middelburg in Mpumalanga southwards to Riversdale in Western Cape and as far west as

Mthatha in Eastern Cape. This species is a small shrublet with tuberous roots and pink to deep

magenta flowers. Variation in P. reniforme in internode length, leaf shape and petiole length

has led to several taxonomic opinions. Ecklon and Zeyher (1835) split P. reniforme into three

distinct species of the genus Cortusina (DC.) Eckl. & Zeyh., namely Cortusina velutina Eckl. &

Zeyh., Cortusina rubro-purpurea Eckl. & Zeyh. and Cortusina reniformis (Curt.) Eckl. & Zeyh., but

these were reduced by Harvey (1860) to varieties as var. velutinum (Eckl. & Zeyh.) Harv., var.

sidaefolium (Thunb.) Harv. and var. reniforme. Knuth (1912) later regarded var. velutinum as a

synonym of P. reniforme and raised var. sidaefolium to species status as P. sidaefolium (Thunb.)

Knuth, which is currently known as P. sidoides DC. In 1995, Dreyer, Marais and Van der Walt

re-established the recognition of infraspecific taxa of P. reniforme by recognising P. reniforme

subsp. velutinum (Eckl. & Zeyh.) Dreyer.

Pelargonium reniforme subsp. reniforme was applied by Dreyer, Marais and Van der Walt (1995) to

plants with elongated internodes (5 mm — 12 mm), reniform leaves and petioles 5 mm — 50 mm

long. This form, although sympatric with P. reniforme subsp. velutinum, is more abundant in the

Port Elizabeth area, Eastern Cape, where it is restricted to drier coastal plains below 300 m.a.s.1.

The form designated as P. reniforme subsp. velutinum has internodes of 1 mm —7 mm in length and

cordate or reniform leaves with petioles of 25 mm — 130 mm in length. Plants with characteristics

corresponding to P. reniforme subsp. velutinum are, according to Dreyer et al. (1995), more

abundant between Grahamstown and Queenstown, Eastern Cape. In all other respects the two

subspecies are identical and their distinction is difficult.

The closely allied Pelargonium sidoides differs from P. reniforme in having black or very dark

purple flowers. It is usually indistinguishable from P. reniforme when not flowering, in having an

http://www.abcjournal.org/ @#¥ge doi:10.4102/abc.v44i1.173_§ —————— = =

- Page 2 of 3

identical habit and leaves that are similar in shape and size,

usually cordate and velvety. The distribution of P. sidoides

overlaps with that of P. reniforme but is wider, the species

occurring throughout Eastern Cape, Lesotho, Free State and

southern Gauteng (Van der Walt & Vorster 1988).

Medicinal use and conservation status

Some Pelargonium species are valued by traditional healers

for their medicinal uses. The roots or extracts of roots of

both P. sidoides and P. reniforme are used to treat diarrhoea,

bronchitis, stomach ailments and respiratory tract infections

(Watt & Breyer-Brandwijk 1962). Both species play an

important role in traditional medicine and are therefore

subject to occasional harvesting for local use, but P. sidoides

is harvested at a much larger scale for export to Europe for

use in the manufacture of a number of medicinal products

(Brendler & Van Wyk 2008).

The conservation statuses of the two subspecies of P. reniforme

are listed as ‘Not Evaluated’ (South African National

Biodiversity Institute [SANBI] 2013a, 2013b), as the difficulty

in distinguishing between them hinders data collection

for conservation assessments. There is concern about the

conservation status of P. reniforme as it can be harvested along

with P. sidoides where the two occur sympatrically because

of their morphological similarity. The conservation status

of P. reniforme is thus currently listed as ‘Near Threatened’ as

a result of suspected widespread overharvesting (Raimondo

et al. 2012). The conservation status of P. sidoides is ‘Least

Concern’ because, although it is heavily harvested, it is a

very widespread and common species (De Castro et al. 2012).

The clarification of morphological differences between the

two subspecies of P. reniforme is regarded as a high priority

by the Threatened Species Programme (D. Raimondo pers.

comm., 29 May 2012) so that the conservation status can be

properly assessed. We therefore investigated the species in

an attempt to resolve the taxonomic problem. Morphological

comparisons were made between the two _ subspecies,

as well as with the nearest relative, P. sidoides, to clarify

taxonomic uncertainty and subsequently contribute to the

conservation of this species. The possibility that variation in

some morphological characters is related to environmental

conditions was also investigated by observing specimens of

both subspecies of P. reniforme in their natural habitat.

Research method and design

Specimens (101 in total) of both subspecies of P. reniforme

from the National Herbarium in Pretoria (PRE), all wild-

collected and covering the entire geographical range of the

subspecies, were studied to assess morphological variation

between the two subspecies. A total of 15 specimens were

previously identified as P. reniforme subsp. reniforme and 86

as P. reniforme subsp. velutinum. In.addition, 140 specimens of

P. sidoides from PRE were investigated in order to determine

whether similar variation occurred in this species. The

diagnostic characters distinguishing the two subspecies — that

is, the lengths of the petioles and the internodes, as well as

the leaf-shapes — were recorded for each specimen. A digital

image of the holotype of P. reniforme subsp. velutinum from

the Herbarium at the Swedish Museum of Natural History,

Stockholm (S) was also examined. Observations of both

subspecies were made in the area between Humansdorp,

East London and Queenstown.

Taxonomic treatment

Pelargonium reniforme Curt. in Curtis’s Botanical Magazine

14: 493 (1800); Pers.: 229 (1806); Desf.: 457 (1809); Willd.: 703

(1809); Ait.: 171 (1812); Haw.: 307 (1812); Sweet: t. 48 (1820);

DC.: 666 (1824); Hoffmg.: 95 (1824); Loudon: 574(1829); Don:

737 (1831); Steud.: 289 (1841); Harv.: 300 (1860); Knuth: 447

(1912); Pole-Evans: 672 (1937); Watt and Breyer-Brandwiik:

454 (1962); Batten and Bokelman: 89 (1966); Smith: 381 (1966);

Clifford: 237 (1970); J.J.A. van der Walt: 40, Figure (1977);

Webb: 69 (1894); Dreyer, Marais and Van der Walt: 325

(1995). Cortusina reniforme (Curt.) Eckl. & Zeyh.: 77 (1835).

Geranium reniforme (Curt.) Andr.: 108 (18000; Poir.: 751 (1812);

Steud.: 679 (1840). Geraniospermum reniforme (Curt.) Kuntze:

95 (1891). Iconotype: Curt.: t. 493 (1800).

Cortusina velutina Eckl. & Zeyh.: 77 (1935). Pelargonium

reniforme var. velutinum (Eckl. & Zeyh.) Harv.: 300 (1860).

Pelargonium reniforme subsp. velutinum (Eckl. & Zeyh.)

Dreyer, in Dreyer, Marais and Van der Walt: 328 (1995).

Type: SOUTH AFRICA. Eastern Cape: ‘campestriis ad

fluvium “Zwartkopsrivier” et collibus in “Adow” [Addo]

(Uitenhage)’, Ecklon & Zeyher 598 (S, holo. — digital image!).

Description

Comparison of diagnostic characters of the two subspecies

Based on field observations as well as morphological

comparison of herbarium specimens, P. reniforme subsp.

velutinum was found to have internodes that were variable

in length and a range of petiole lengths overlapping with

those of subsp. reniforme (Table 1). This finding contradicts

the descriptions given by Dreyer ef al. (1995) to distinguish

the two subspecies.

TABLE 1: Summary of diagnostic characteristics of Pelargonium species recorded from herbarium specimens.

Character

Leaf shape Reniform

Petiole length measurements from herbarium specimens (mm) 10-115

Petiole length according to Dreyer et al. (1995) (mm) 5-508

Internode length (mm) 1-15

Internode length according to Dreyer et al. (1995) (mm) 5-12

Pelargonium reniforme subsp. reniforme

Pelargonium reniforme subsp. velutinum Pelargonium sidoides

Cordate or reniform Cordate or reniform

6-138 12-200

25-130 Not given

1-10 1-3

1—/ Not given

Source: Comparative data where stated from Dreyer, L.L., Marais, E.M. & Van der Walt, J.J.A., 1995, ‘A subspecific division of Pelargonium reniforme Curt. (Geraniaceae)’, South African Journal of

Botany 61(6), 325—330

http://www.abcjournal.org

dot:10.4102/abc.v44i1.173

Character variation was plotted on a scatter plot chart

(Figure 1), which shows that all variation overlaps in both

subspecies. This implies that these characters cannot be

used to distinguish the two subspecies. The petiole length

of subspecies velutinum overlaps completely with that of

subspecies reniforme and this characteristic is therefore

also not useful to distinguish the two subspecies. Most of

the specimens with characteristics that correspond with

subspecies velutinum have cordate leaves; although, there

are a few that are reniform. All specimens corresponding to

subspecies reniforme have reniform leaves. Figure 1 shows

that variation in the leaf shape is not correlated with petiole

length or internode length and therefore this character is too

variable to reliably be used to distinguish subspecies.

Pelargonium sidoides is vegetatively similar to the descriptions

given in Dreyer et al. (1995) for subspecies of P. reniforme;

however, one consistent characteristic of P. sidoides is the

short internodes (usually 1 mm but never longer than 3 mm

in length), whereas the internode length in P. reniforme varies

between specimens. The other consistent difference between

the two taxa is the flower colour, as described above.

Influence of habitat on morphology

Although specimens with the form P. reniforme subsp.

reniforme are concentrated around Port Elizabeth,

observations of plant characteristics in this area revealed

that a possible cause for variation is the microhabitat in

which the plant was growing. Plants growing in the shade

of dense grass in the Port Elizabeth area tended to have

both longer internodes (as in subsp. reniforme), as well

as longer petioles (as in subsp. velutinum). It is possible

that availability of nutrients, water and sunlight affects

internode and petiole lengths, which could explain much of

the variability of these characteristics in P. reniforme.

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Series 1, Pelargonium reniforme subsp. velutinum with cordate leaves; Series 2, Pelargonium

reniforme subsp. velutinum with reniform leaves; Series 3, Pelargonium reniforme subsp.

reniforme with reniform leaves.

FIGURE 1: Scatter-plot chart of character variation within Pelargonium reniforme

subsp. reniforme and Pelargonium reniforme subsp. velutinum.

http://www.abcjournal.org/ 29 | doi:10.4102/abce.v44i1.173

Short Note

Conclusion

The findings of this study show that it is not possible to

reliably distinguish between P. reniforme subsp. reniforme

and P. reniforme subsp. velutinum and we recognise a single

variable species for all populations.

Acknowledgements

The Botanical Education Trust is gratefully acknowledged

for the funding received to enable this project to be carried

out. We are grateful to Dr John Manning and Mr Tony Dold

for discussions and comments on the manuscript. Two

anonymous reviewers are gratefully acknowledged for

improvements to the manuscript.

Competing interests

The authors declare that they have no financial or personal

relationships that may have inappropriately influenced them

in writing this article.

Authors’ contributions

M.A. (SANBI) examined herbarium specimens and took

measurements under the training and guidance of J.E.V.

(SANBI). M.A. contributed to writing of the manuscript

and collated background information. J.E.V. undertook

fieldwork, contributed to writing the manuscript and editing

it and supervised the project.

References

Bakker, FT., Culham, A., Hettiarachi, P., Touloumenidou, T. & Gibby, M., 2004,

‘Phylogeny of Pelargonium (Geraniaceae) based on DNA sequences from three

genomes’, Taxon 53, 17-28.

Brendler, T. & Van Wyk, B.-E., 2008, ‘A historical, scientific and commercial perspective

on the medicinal use of Pelargonium sidoides (Geraniaceae)’, Journal of

Ethnopharmacology 119(3), 420-433.

De Castro, A., Vlok, J.H., Newton, D., Motjotji, L. & Raimondo, D., 2012, ‘Pelargonium

sidoides DC’, National assessment: Red list of South African plants, version

2013.1, viewed 12 November 2013, from http://redlist.sanbi.org/species.

php ?species=1976-307

Dreyer, L.L., Marais, EM. & Van der Walt, J.J.A., 1995, ‘A subspecific division of

Pelargonium reniforme Curt. (Geraniaceae)’, South African Journal of Botany

61(6), 325-330.

Ecklon, C.F. & Zeyher, K.L.P., 1835, Enumeratio plantarum africae australis extratropicae

quae collectae, determinatae et expositae, Perthes & Besser, Hamburg.

Harvey, W.H., 1860, ‘Geraniaceae’, in W.H. Harvey & O.W. Sonder, Flora capensis,

vol. 1, p. 129, Hodges Smith, Dublin.

Knuth, R., 1912, ‘Geraniaceae’, in H.G.A. Engler (ed.), Das Pflanzenreich, vol. 4, p. 129,

Wilhelm Engelmann, Berlin.

Raimondo, D., Victor, J.E., Dold, A.P. & De Castro, A. 2012, ‘Pelargonium reniforme

Curtis’, National assessment: Red list of South African plants, version

2013.1, viewed 12 November 2013, from http://redlist.sanbi.org/species.

php ?species=1976-4012

Roux, J.P., 2013, ‘Validating Pelargonium sect. Reniformia’, Bothalia 43(2), 233.

South African National Biodiversity Institute, 2013a, ‘Pelargonium reniforme Curtis

subsp. reniforme’, National assessment: Red list of South African plants, version

2013.1, viewed 12 November 2013, from _http://redlist.sanbi.org/species.

php ?species=1976-284

South African National Biodiversity Institute, 2013b, ‘Pelargonium reniforme Curtis

subsp. velutinum (Eckl. & Zeyh.) Dreyer’, National assessment: Red list of South

African plants, version 2013.1, viewed 12 November 2013, from http://redlist.

sanbi.org/species.php ?species=1976-363

Van der Walt, J.J.A., 1985, ‘A taxonomic revision of the type section of Pelargonium

UHerit. (Geraniaceae)’, Bothalia 15(3&4), 345-385.

Van der Walt, J.J.A. & Vorster, P.J., 1988, Pelargoniums of southern Africa, vol. 3,

National Botanic Gardens, Kirstenbosch.

Vorster, P., 2000, ‘Geraniaceae’, in P. Goldblatt & J.C. Manning (eds.), ‘Cape plants:

A conspectus of the Cape Flora’, Strelitzia 9, 516-528, National Botanical Institute,

Cape Town.

Watt, J.M. & Breyer-Brandwijk, M.G., 1962, Medicinal and poisonous plants of southern

and eastern Africa, 2nd edn., pp. 454-455, E&S Livingstone Ltd, Edinburgh.

Page 1 of 4

Ledebouria caesiomontana A.jJ.Hankey & N.Hahn

(Hyacinthaceae: Hyacinthoideae): A new species from

the Blouberg centre of endemism, Limpopo, South Africa

Authors:

Andrew J. Hankey*

Norbert Hahn?

Matt H. Buys**

Affiliations:

1South African National

Biodiversity Institute, Walter

Sisulu National Botanical

Garden, South Africa

*Herbarium

Soutpansbergensis,

South Africa

*National Forestry

Herbarium, New Zealand

Forest Research Institute,

New Zealand

“Department of Botany

and Zoology, University of

Stellenbosch, South Africa

Correspondence to:

Andrew Hankey

Email:

a.hankey@sanbi.org.za

Postal address:

PO Box 2194, Wilro Park

1731, South Africa

Dates:

Received: 06 Feb. 2014

Accepted: 31 July 2014

Published: 04 Nov. 2014

How to cite this article:

Hankey, A.J., Hahn,

N. & Buys, M.H.,

2014, ‘Ledebouria

caesiomontana A.J.Hankey

& N.Hahn (Hyacinthaceae:

Hyacinthoideae): A new

species from the Blouberg

centre of endemism,

Limpopo, South Africa’,

Bothalia 44(1), Art. #119,

4 pages. http://dx.doi.

org/10.4102/abc.v44i1.119

Read online:

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[ltr cer toread online.

Background: Ledebouria Roth (Hyacinthaceae: Hyacinthoideae) is a largely African genus with,

more or less, 40 species occurring in South Africa. The species was first collected in December

1990 by R. Archer [Archer, R.H. 503 (PRE)] and remained un-identified. Subsequently, N. Hahn

also collected the species in 1992 [N. Hahn 444 (ZPB)] and, upon failing to resolve the identity

of the specimen, he approached A. Hankey who initiated further collaborative research.

Objective: To describe the new species of Ledebouria from Blouberg mountain massif in

Limpopo Province, South Africa.

Method: Relevant existing specimens in herbaria were examined and morphological

characters and states noted. The type specimen was collected during an expedition under the

guidance of Prof. Dirk Bellstedt accompanied by Mr Adam Harrower.

Results: Ledebouria caesiomontana A.J. Hankey & N.Hahn sp. nov. was described and illustrated.

The new species was distinguished from its closest relative, Ledebouria papillata S.Venter, by

the ovary which lacks basal lobes, as well the absence of cataphylls and the irregular papillate

ridges present only on the upper leaf surfaces.

Conclusion: Ledebouria caesiomontana is a new species restricted to the Blouberg mountain

massif in Limpopo Province, South Africa. Initial estimates deem the species to be vulnerable

(VU D2) as a result of especially anthropogenic-induced disturbances on the Blouberg.

Introduction

The genus Ledebouria Roth (Hyacinthaceae: Hyacinthoideae) was first erected by Roth (1821)

when he described Ledebouria hyacinthina Roth from India. Baker (1870) subsequently placed

the genus under Scilla L. where it typified section Ledebouria. Jessop (1970) resurrected the

genus and recognised 15 South African species, characterised by the possession of bulbs with

deciduous leaves and erect to mostly somewhat flaccid inflorescences with basal ovules paired

in each locule, together with a stipitate ovary. The most recent revision of Ledebouria by Venter

(1993) was followed by a synopsis of the genus by the same author (Venter 2008), wherein he

recognises 39 South African species and makes reference to more than 60 species occurring in

sub-Saharan Africa, with one or two each in India and Madagascar. Despite the large number

of species currently recognised in Ledebouria, no formal infra-generic classification exists to date.

This study was initiated by N. Hahn, who collected the species from Blouberg mountain massif

in Limpopo Province, South Africa, in 1992 [N. Hahn 444 (ZPB)] and, upon failing to resolve

the identity of the specimen, approached the first author for assistance. Subsequently, an

examination of existing herbarium specimens revealed un-identified specimens that belonged to

the same entity. Recollections by some of the authors, coupled with an analysis of characters and

states based on herbarium specimens, fieldwork and living material, led to the conclusion that a

new species was in hand. In this article, we describe the new species of Ledebouria.

Research method and design

Relevant specimens in the South African National Herbarium (PRE) and the Herbarium

Soutpansbergensis (ZPB) formed the bulk of the material available for study. In addition, a

collection of living plants housed at the Walter Sisulu National Botanical Garden, Roodepoort,

Gauteng Province, provided material for illustrative and research purposes. The unknown entity

was compared on a macromorphological and micromorphological basis with similar looking

species. Measurement of plant parts was performed by hand using a combination of rulers and

callipers.

doi:10.4102/abc.v44i1.119

pape Ona

Taxonomic treatment

Ledebouria caesiomontana A.J.Hankey &

N.Hahn sp. nov.

Type: SOUTH AFRICA. Limpopo: 2329 (Blouberg): Beauley,

23.08467 °S, 29.00076 °E, 1658 m.a.s.l. cliff ledges in forest,

(-AA), 24 Mar. 2007, Hankey, A.J. 2129 (PRE, holo; NBG, iso).

Description

Plants mostly solitary to weakly gregarious by sobolifery

(proliferating from the base of the bulb). Bulbs: hypogeal

20 mm — 25 mm x 10 mm — 15 mm, ovoid, dead bulb

scales membranous, pale brown, live bulb scales tightly

arranged, whitish, fleshy, truncate apically, with threads

when torn. Leaves: (3—-)4(-5), dull green, linear-oblanceolate,

70 mm — 100 mm x 10 mm — 15 mm, synanthous, apex acute,

base canaliculate, with threads when torn, spreading, green,

with irregular longitudinal papillate ridges, maculate with

dark green to purple spots and blotches adaxially, glabrous,

wholly purple-red, or green suffused with purple-red

abaxially, margin entire, minutely papillate. Inflorescence:

one per bulb, erect, 65 mm — 120 mm long, raceme

30 mm — 40 mm long, conical, with 12-20 loosely arranged

flowers, scape terete, glabrous, 50 mm — 80 mm long, reddish-

green. Bracts: present, 1.0 mm x 0.5 mm, deltoid, fleshy,

reddish-green, prophylls absent. Pedicels: 5 mm — 7 mm long,

pink. Perianth: campanulate; tepals oblong, cucullate apically,

reflexed, 5.0 mm — 6.0 mm x 1.8 mm, pink with greenish-

brown vitta. Stamens: epitepalous, erect, filaments 6 mm long,

filiform, violet; anthers yellow, 0.5 mm long, dorsifixed. Ovary:

six lobed, greenish, depressed ovoid, 1 mm x 2 mm, stipitate,

stipe 0.5 mm x 0.5 mm, basal lobes absent, style 5 mm long,

violet. Flowering time: October to December (Figure 1).

Distribution and ecology

Ledebouria caesiomontana occurs in Northern Mistbelt Forest

vegetation (Mucina & Rutherford 2006) on the Blouberg

massif. Specifically, it has been recorded from low deciduous

forest and forest margins in shallow, moss-covered lithosols

and rock crevices under the shade of woody vegetation. The

species is cryptic and easily overlooked in its habitat because

the bulbs are often tightly wedged in narrow rock crevices.

This species is thus far only known from the Blouberg

massif, where it has only been recorded from five collections

(Figure 2).

The description of this new species brings the total of

Blouberg endemic plant species to four (Hahn 2006), with the

following endemic species previously known being: Berkheya

radyeri Roessler, Cineraria cyanomontana Cron (Asteraceae)

and Streptocarpus longiflorus (Hilliard & B.L.Burtt)

T.J.Edwards (Gesneriaceae). The new species has thus far

not been found in the Soutpansberg, an area known to share

13 near endemic taxa with the Blouberg (Hahn 2006).

http://www.abcjournal.org/ ER

Short Note

Etymology

The specific epithet caesiomontana alludes to the Blouberg

(meaning Blue Mountain) in the Limpopo Province of South

Africa, from which the new species was collected.

Conservation status

Owing to L. caesiomontana’s cryptic nature, it may easily be

overlooked in the field and this may explain the low numbers

of collections. Ledebouria caesiomontana is uncommon in its

known distribution range but more extensive fieldwork may

reveal additional populations.

The forests on the Blouberg are systematically being

eradicated as a result of slash and burn practices. To date,

no official study has been undertaken on the Blouberg to

ascertain the extent these activities could potentially have

on the conservation of L. caesiomontana or the Blouberg as

a whole. An urgent study needs to be conducted on the

Blouberg to assess the impact and inform the conservation

measures required to halt the rapid loss of forests in the

area. In the light of the preceding factors, we expect that

the species would be best ascribed as VU D2 in terms of the

International Union for Conservation of Nature’s (IUCN)

Red List status (IUCN 2012).

Diagnosis

Ledebouria caesiomontana is most similar to Ledebouria

papillata, from which it is distinguished by several

characters (Table 1). The new species lacks the

characteristic basal ovary lobes of L. papillata — not

illustrated by Venter (1993, 2008) but clearly noted in the

descriptions. Furthermore, L. caesiomontana differs in the

absence of regular longitudinal rows of papillae on the

scape and leaves, instead having irregular papillate ridges

only on the adaxial surface of the leaf and not on the scape

or the abaxial leaf surface. This species also lacks the two

basal cataphylls noted by Venter (2008) as diagnostic for

L. papillata.

Ledebouria asperifolia (Van der Merwe) S.Venter (Venter2008)

also possesses longitudinal rows of papillae on the lower

leaf surface (and occasionally on the upper leaf surfaces) and

may be superficially similar to L. caesiomontana. However,

L. asperifolia is larger in all respects, and has purplish-brown

dead bulb scales persistent on the bulb. The distribution of

these two species is furthermore, distinct (Figure 2).

Additional specimens examined

SOUTH AFRICA. Limpopo: 2329 (Blouberg): Blouberg,

Blouberg Nature Reserve, 13 Dec. 1990, Venter, S. 13507

(PRE); Blouberg NR, Farm Dantsig 3, 05 Dec. 1990, Archer,

R.H. 503 (PRE); Blouberg, Beauley, 23.0776 °S, 28.99324 °E,

1692 m.a.s.l., cliff ledges in forest, 24 Mar. 2007, Hankey,

A.J. 2130 (PRE) (-AA). 2328 (Tolwe): Blouberg, Beauley,

23°4’40.14” S , 28°59’38.50” E, 1710 m.a.s.1., in flower, 09 Dec.

1992, N. Hahn 444, (ZPB) (—BB).

dai A07

Page 3 of 4 Ee Short Note

Source: \\lustrations by S. Burrows; specimen voucher: Hankey, A.J. 2129

FIGURE 1: Illustrations of Ledebouria caesiomontana depicting, (a) flowering plant, (b) adaxial leaf surface showing ridges and ornamentation detail, (c) inner and outer

tepal section showing stamens, (d) flower, pedicel and floral bract, (e) ovary lateral and dorsal view and (f) flower, section showing ovary and stamens.

ee itty / WW WiddcjOunnalonrey, 32 | doi:10.4102/abc.v44i1.119

TABLE 1: Differences between Ledebouria papillata and Ledebouria caesiomontana.

Page 4 of 4 — SONG CMe

Ledebouria caesiomontana

Category Ledebouria papillata (Venter 2008)

Distribution Pietersburg plateau, Gauteng and Eastern Cape Provinces

Bulb With prominent neck and 2 basal cataphylls reaching ground level — not

shown in illustration by Venter (2008)

Leaves 2-4, glossy dark green, with venation prominent

Leaf base Sub-petiolate with vertical purple stripes

Epidermal ornamentation

scape

Inflorescence Erect

Perianth 2.0 mm — 3.5 mm long

Ovary Basal lobes present — not shown in illustration by Venter (2008)

23°0'0"S

24°0'0"S

25°0'0"S

26°0'0"S

} Legend

i BHO Ors Ledebouria

caesiomontana

Ledebouria

papillata

Altitude

- High

ie ,..,

28°0'0"S

120 180

28°0'0"E

29°0'0"E

27°0'0"E

30°0'0"E

31°0'0"E

Source: Map drawn by M. Lotter

FIGURE 2: Known distribution of Ledebouria caesiomontana and Ledebouria

papillata.

Acknowledgements

Hugh Glen is thanked for comments on the specific epithet,

Sandie Burrows for the illustration and Mervyn Lotter for

the distribution map. The local Tribal Authority is thanked

for allowing access to the property.

Longitudinal rows of papillae on adaxial and abaxial leaf surfaces and

Endemic to the Blouberg, Limpopo Province

Without prominent neck, cataphylls absent

3-5, dull mid-green, with venation obscure

Clasping, canaliculate without vertical stripes

Irregular longitudinal ridges on the adaxial leaf surface only; Abaxial leaf

surface and scape glabrous

Spreading, becoming flaccid

5.0 mm—6.0 mm long

Basal lobes absent

Source: Ledebouria papillata data taken from Venter, S., 2008, ‘Synopsis of the genus Ledebouria Roth (Hyacinthaceae) in South Africa’, Herbertia 62, 85-155

Competing interests

The authors declare that they have no financial or personal

relationships that may have inappropriately influenced

them in writing this article.

Authors’ contributions

A.J.H. (Walter Sisulu National Botanical Garden) was

responsible for the morphological study and wrote the

description, whilst N.H. (Herbarium Soutpansbergensis)

provided the conservation, ecological and phytogeographic

notes. M.H.B. (New Zealand Forest Research Institute)

undertook a supervisory and advisory role, providing

taxonomic guidance and editing.

References

Baker, J.G., 1870, ‘Monograph of Scilla: Sections Ledebouria and Drimiopsis’,

Saunder's Refugium Botanicum 3, 1-18.

Hahn, N., 2006, ‘Floristic diversity of the Soutpansberg, Limpopo Province, South

Africa’, unpublished PhD thesis, Department of Plant Science, University of

Pretoria.

International Union for Conservation of Nature (IUCN), 2012, [UCN Red List categories

and criteria, version 3.1, 2nd edn., |UCN, Gland.

Jessop, J.P., 1970, ‘Studies in the bulbous Liliaceae: 1. Scilla, Schizocarphus and

Ledebouria’, Journal of South African Botany 36, 233-266.

Mucina, L. & Rutherford, M.C., 2006, ‘The vegetation of South Africa, Lesotho and

Swaziland’, Strelitzia 19, South African National Biodiversity Institute, Pretoria.

Roth, A.G., 1821, ‘Ledebouria hyacinthina’, in Novae Plantarum species Indiae

Orientalis, Ex collectione doct. Benj. Heynii: cum descriptionibus et

observationibus, pp. 195-196, Sumptibus H. Vogleri, Halberstadii. http://dx.doi.

org/10.5962/bhi.title.10723

Venter, S., 1993, ‘Revision of the genus Ledebouria Roth (Hyacinthaceae) in

South Africa’, unpublished MSc thesis, Department of Botany, University of

Natal.

Venter, S., 2008, ‘Synopsis of the genus Ledebouria Roth (Hyacinthaceae) in South

Africa’, Herbertia 62, 85-155.

http://www.abcjournal.org/ 33 | doi:10.4102/abe.v44i1.119

S02 ———

Gladiolus filiformis, a poorly known species from

Authors:

Norbert Hahn!

Hermien Roux?

Affiliations:

‘Department of Economic

Development, Environment,

Conservation and Tourism,

North West Provincial

Government, South Africa

Correspondence to:

Norbert Hahn

Email:

nhahn@soutpansberg.com

Postal address:

Private Bag X2039,

Mmabatho 2735,

South Africa

Dates

Received: 25 June 2013

Accepted: 02 July 2014

Published: 04 Nov. 2014

How to cite this article:

Hahn, N. & Roux, H., 2014,

‘Gladiolus filiformis, a poorly

known species from North

West Province, South Africa’,

Bothalia 44(1), Art. #174,

4 pages. http://dx.doi.

org/10.4102/abc.v44i1.174

Licensee: AOSIS

OpenJournals. This work

is licensed under the

Creative Commons

Attribution License.

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North West Province, South Africa

Background: Gladiolus filiformis was described in 1998 from only the type specimen, which

was collected in 1977.

Objectives: The aim of this article was to expand our knowledge of the morphology,

distribution, ecology, and conservation status of this poorly known species.

Method: Data presented were based on field observations, with dissections and diagnostic

descriptions conducted in the field.

Results: A comprehensive account of the morphology, distribution, ecology and conservation

status for G. filiformis was presented for the first time.

Conclusion: Gladiolus filiformis is more common than previously thought. The species is not

an edaphic specialist and it occurs mainly on sediments of the Transvaal supergroup. Its

heteromorphically winged fruit were described here, showing an intermediate form between

Gladiolus pretoriensis and Gladiolus oatesii.

Introduction

The North West Department of Economic Development, Environment, Conservation and Tourism

is presently conducting a biodiversity inventory of the rare, endangered and endemic plants in the

province, including Gladiolus filiformis, which is considered to be rare and endangered.

In December 2010, G. filiformis Goldblatt & J.C. Manning was rediscovered by the authors.

This species was previously known only from its type locality and was described from a single

herbarium specimen (Goldblatt & Manning 1998).

The aim of this article is to expand our knowledge of the morphology, distribution, ecology and

conservation status of this poorly known species.

Research method and design

All presently known populations of G. filiformis were surveyed over a period of three flowering

seasons, from December 2010 to February 2013. Approximately 300 live plants were examined.

Live plants were measured and dissected in the field. Distribution and ecological data were

captured with the aid of a Trimble Juno ST, Personal Digital Assistant. Co-ordinates were

recorded using datum WGS 84. All data gathered from the biological surveys are stored in a

Global Information System database to be incorporated into the planned Biodiversity Information

Management System for North West Province.

Landowners’ permission was obtained for this study and no permits were required. Specimens

were deposited in the Compton Herbarium (NBG) and Herbarium Soutpansbergensis (ZPB).

Taxonomic treatment

Gladiolus filiformis Goldblatt & J.C. Manning in Gladiolus South Africa:

182 (1998).

Type: SOUTH AFRICA: ‘North Eastern Cape’ [North West Province]: [2525: Mafikeng], Gopane

Mountains, rocky koppie, rocky red soil, altitude 1100 m.a.s.l. [sic], [-BD)], 29 Dec. -1977,

C.P. Peeters, N.P. Gericke & G.G. Burelli 509 (PRE, holo.!).

Description

Plants: 350 mm = 500(—720) mm tall. Corm: obconic, up to 30 mm in diameter, ivory coloured; tunics

of reticulate fibres. Cataphylls: brown, membranous, protruding about 20 mm above ground. Leaves:

§ doi:10.4102/abc.v44i1.174

usually three, occasionally four, basal leaves reaching to base

of inflorescence, second leaf sheathing lower third of stem,

blades terete and four-grooved with margins and veins

thickened, 1.0 mm — 2.5 mm in diameter. Stems: erect, usually

simple, up to 2 mm in diameter. Inflorescence: a (5—)6 mm —

10(-12) mm flowered spike. Bracts: green tinged, reddish at

apex, drying after anthesis, outer bracts 9.3 mm — 24.4 mm x

3.8 mm — 7.0 mm long, inner bracts 0.833-1.000 times shorter

than outer bracts. Flowers: usually light mauve, rarely tinged

pink, dorsal tepal usually dark lilac with burgundy markings

at base; upper lateral tepals midrib burgundy on the inside;

lower lateral tepals with various white and darker markings,

apex with two burgundy lines on outside; lower median tepal

with or without various white and darker markings, base

with single or double burgundy line on the inside; unscented

by day. Length of flower measured from base of tube to tip of

dorsal tepal (49.2-)51.2 mm — 64.0(—70.0) mm long; perianth

tube filiform, curved, (25.4-)27.1 mm — 36.5(-40.8) mm long;

dorsal tepal elliptic to broadly elliptic (19.8-)20.3mm —

24.1 (—24.8) mm x (14.7—)14.9 mm — 19.3(—22.0) mm, apex acute,

base tapering; upper lateral tepals elliptic (15.1-)15.9 mm —

Daleo\(—2'520)) sam (ON N=)9)7 som — 19 (E136) mim,

apex acuminate, base tapering; lower lateral tepals elliptic

to narrowly elliptic (14.6-)154mm — 18.3(-20.3)mm x

(5.8-)5.9 mm — 8.8(—-10.6) mm, apex acuminate, base attenuate;

lower median tepal elliptic (14.6—)15.4 mm — 18.3(-20.3) mm x

(6.5-)7.3 mm —10.3(-11.7) mm, apex acuminate, base tapering.

Stamens: outer filaments from ovary (31.7-)32.mm —

42.1(-45.5) mm long, unattached section (9.2—)10.3 mm — 13.0

(-14.2) mm long; median filament from ovary (32.5-)33.5 mm

— 43.5(-47.3) mm long, unattached section (8.6—)10.6 mm —

13.9 mm long; anthers (5.3-)5.9 mm — 8.1(-9.0) mm long,

yellow. Ovary: ovoid (2.5—)2.9 mm — 4.2(-4.4) mm long; style

(31.8-)37.7 mm — 49.8(-54.2) mm long, branching at apex,

branches (2.8-) 3.0 mm — 4.6(—6.0) mm long. Capsule: ellipsoid,

7.7 mm — 11.8 mm long. Seeds: irregularly shaped, 1.8 mm —

3.1mm long, brown, surface granulate, wings heteromorphous

(wings of different shapes), apical wing 0.9 mm — 2.4 mm

long; basal wing 0.2 mm — 0.7 mm long, lateral wing 0.2 mm —

0.5 mm long with the opposing wing usually absent or

underdeveloped. Flowering period: Mainly December, but can

extend from October to February (Figure 1).

The north-western plants, including the type, are smaller than

those found to the south-east. The inflorescences of the type

specimen have only two flowers but the plant was described

as having five flowers. Five flower inflorescences are the

lower limit for this species, with the number usually lying

between six and ten, with an upper limit of twelve observed.

The flowering structure of the type specimen is also smaller

than those observed. The perianth tube of the type specimen

was stated as approximately 20 mm, whereas the length in

the plants studied was found to be between 27.1 mm and

36.5 mm, with the lower and upper limit between 25.4 mm

and 40.8 mm. The shape, length and colouration of the dorsal

tepals, upper tepals, lower lateral tepals and lower median

tepals are described for the first time as they were omitted

in the original description of G. filiformis. The rare pink form

of G. filiformis has not been recorded before. The unattached

Short Note

outer and median filament sections were found to be of

different sizes with the latter usually shorter; this suggests

a possible diagnostic feature. No fruiting characteristics

were recorded for the species prior to this study. The newly

described heteromorphically winged seeds of G. filiformis,

with the lateral wings absent or underdeveloped and the

apical wing up to 2.4 mm long, are intermediate between the

wingless seeds of Gladiolus pretoriensis and the broad, evenly

winged seeds of Gladiolus oatesit, which are up to 5 mm long

(Lewis, Obermeyer & Barnard 1972).

Related taxa

Gladiolus filiformis has an elongated, recurved, filiform

perianth tube, which is diagnostic of the species (Goldblatt

& Manning 1998). Gladiolus pretoriensis Kuntze is similar and

is also a Southern Bankenveld endemic, but it is usually a

smaller plant with smaller pink flowers. Gladiolus pretoriensis

fruits are said to be wingless, whilst those of G. filiformis are

heteromorphically winged. Gladiolus filiformis could also be

confused with G. oatesii Rolfe, but this species has a funnel-

shaped perianth tube and linear, as opposed to lanceolate,

leaves.

Distribution and ecology

The type locality is stated as Gopane Mountains on a

rocky koppie, growing in rocky red soil at an altitude of

1100 m.a.s.l. The Gopane Mountains are not indicated

on any map, but two towns with the name are. The first

reference applies to the remains of an old military base east

of Skilpadhek Border Post (2525BC). The whole area around

the base is above 1220 m.a.s.l. and we could find no rocky

koppie with red soil in the vicinity. The second Gopane is

the town in which the Livingston Mission Station (2525BD)

is situated. Most of the area is above 1180 m.a.s.1. and there is

a rocky koppie to the north-west, with red soil. This koppie

is known as Sedukwane (1303 m.a.s.1.). It was on this koppie

that we located G. filiformis at an altitude of 1290 m.a.s.L,

hence this is most likely the type locality (Figure 2). Peeters,

Gericke and Burelli, who collected the type, are therefore

most probably in error as it is impossible for G. filiformis to

have been collected at an altitude of 1100 m.a.s.1.

Gladiolus filiformis is presently only known from the extreme

western sections of the Southern Bankenveld (Partridge

et al. 2010), where it grows on hills, hill slopes and plateaus.

The plant is not an edaphic specialist, having been observed

growing in soils derived from sediments of the Transvaal

supergroup, comprising banded ironstone, conglomerate

and chert or dolomite areas. The type locality is situated

on a dolerite koppie, the remains of a post-Transvaal

supergroup volcanic plug. Most populations of G. filiformis

occur in grassland, except for those towards its western

extremities, including the type locality, which are found in

open scrubland. Its presently known distribution, from the

north-west (Maphephane) to its south-eastern extremity

(Tweefontein), stretches across an expanse of 40 km.

Gladiolus filiformis was found growing sympatrically with

Gladiolus permeabilis Delaroche subsp. edulis (Burchell ex

http://www.abcjournal.org/ 35 | doi10:4102/abev44i.174 —— SS

by Hermien Roux

FIGURE 1: Gladiolus filiformis vegetative and reproductive morphology depicting, (a) inflorescence, (b) capsules — open and unopened and (c) seed.

Scale bar: (a) 4.0 mm; (b) 3.6 mm and (c) 0.5 mm

Source: \\lustrations

South Africa

4 Gladiolus filiformis

25°45’E 26°0'E

Source: Figure provided by Norbert Hahn

FIGURE 2: Distribution of Gladiolus filiformis, (a) regionally within South Africa and (b) locally within North West Province.

Ker Gawler) Obermeyer, but not with its near relatives

G. pretoriensis or G. oatesit.

Conservation status

A word of caution is necessary in attempting a conservation

assessment of this species. Flowering of the species was

prolific in the seasons of December 2010 and December 2012

— January 2013 as a result of good rains in the beginning of

December in both seasons, with plants flowering in their

thousands. The period of October 2011 — February 2012 was a

poor rainfall season, with only a few plants flowering. In the

good seasons the plants bloomed mainly from December to

January, whilst in the poor times they flowered sporadically

from October to February.

At the type locality, grazing by goats could pose a threat,

a road runs through two of the southern populations, a

radio station is built at the third population and a pipeline

transects the eastern populations. Notwithstanding the

above there are no indications of the populations declining.

The present known extent of occurrence of this species

is 287 km’, thus it does not qualify as ‘Critically Rare’

(Goldblatt & Victor 2009). The estimated number of mature

individuals far exceeds 100 000 and the populations seem

stable, thus excluding criteria VU C or VU D1 (International

Union for Conservation of Nature 2001). A status of ‘Least

Concern’ is proposed, but this will need to be revised if

future mining activities are embarked upon in the area.

Additional collections

BOTSWANA: Troverston, 25°14'30.024"S, 25°44’07.104" E,

11 Jan. 2013, N. Hahn 3038 (NBG, ZPB). SOUTH AFRICA:

North West Province: Sedukwana, 25°17'59.520"S,

25°47'55.038” E; 25°28'44.538" S, 25°50'12.276" E,

15 Dec. 2010, N. Hahn 2767, 2772 (NBG, ZPB); Moilwas,

http://www.abcjournal.org/ @eyé

25°26'37.992" S, 25°49'11.016” E, 16 Feb. 2013, N. Hahn,

3053 (ZPB); Tweefontein, 25°32'53.052" S, 25°55’42.618”" E,

01 Jan. 2013, N. Hahn 3033 (NBG, ZPB).

Acknowledgements

Dr John Manning is thanked for his help in identifying

Gladiolus material. The Directorate Environmental Planning

and Coordination, Department of Economic Development,

Conservation and Tourism, North West Provincial

Government, Republic of South Africa is thanked for making

the opportunity available to the senior author to study

G. filiformis as part of his job description.

Competing interests

The authors declare that they have no financial or personal

relationships that may have inappropriately influenced

them in writing this article.

Authors’ contributions

H.R. (North West Provincial Government) assisted with

fieldwork and data capture, helped write the manuscript and

drew the accompanying plate. N.H. (North West Provincial

Government) did most of the fieldwork, the dissections for

the diagnostic description and wrote most of the article.

References

Goldblatt, P. & Manning, J.C., 1998, Gladiolus in southern Africa, Fernwood Press,

Vlaeberg.

Goldblatt, P. & Victor, J.E., 2009, ‘Gladiolus filiformis’, in D. Raimondo, L. von Staden,

W. Foden, J.E. Victor, N.A. Helme, R.C. Turner et al. (eds.), ‘Red list of South

African plants 2009’, Strelitzia 25, p. 146, South African National Biodiversity

Institute, Pretoria.

International Union for Conservation of Nature, 2001, /UCN red list categories and

criteria: Version 3.1, \UCN Species Survival Commission, Gland, Switzerland.

Lewis, G.J., Obermeyer, A.A. & Barnard, T-T., 1972, ‘Gladiolus: A revision of the South

African species’, Journal of South African Botany Suppl. 10.

Partridge, T.C., Dollar, E.S.J., Moolman, J. & Dollar, L.H., 2010, ‘The geomorphic

provinces of South Africa, Lesotho and Swaziland: A physiographic subdivision for

earth and environmental scientists’, Transactions of the Royal Society of South

Africa 65(1), 1-47. http://dx.doi.org/10.1080/00359191003652033

doi:10.4102/abc.v44i1.174

Page 1 of 4

Lectotypification of Kniphofia pauciflora Baker

Authors:

Himansu Baijnath?

Syd Ramdhani?

Affiliations:

'School of Life Sciences,

University of KwaZulu-Natal,

Westville Campus,

South Africa

Correspondence to:

Himansu Baijnath

Email:

baijnathh@ukzn.ac.za

Postal address:

Private Bag X54001, Durban

4000, South Africa

Dates:

Received: 22 Mar. 2013

Accepted: 10 Feb. 2014

Published: 12 Nov. 2014

How to cite this article:

Baijnath, H. & Ramdhani,

S., 2014, ‘Lectotypification

of Kniphofia pauciflora

Baker (Asphodelaceae:

Asphodeloideae)’, Bothalia

44(1), Art. #151, 4 pages.

http://dx.doi.org/10.4102/

abc.v44i1.151

Licensee: AOSIS

OpenJournals. This work

is licensed under the

Creative Commons

Attribution License.

Read online:

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(Asphodelaceae: Asphodeloideae)

Background: Problems amongst the syntypes and duplicates were noted for the critically

endangered Kniphofia pauciflora Baker.

Objective: To resolve the typification of K. pauciflora.

Method: Literature and specimens (including type material) were examined from relevant

herbaria.

Results: Lectotypification was necessary in this case. The lectotype designated was Wood

1096 (K) from ‘Natal, Claremont flat’ [KwaZulu-Natal, Clairmont flat]. This was undertaken

to specify a single type specimen and to clarify the status of the duplicates. Also included

is an account of the last remaining population of K. pauciflora at the Clairwood Racecourse,

Durban, South Africa.

Conclusion: The lectotype was designated as Wood 1096 (K).

Introduction

The earliest herbarium collection of Kniphofia pauciflora Baker that we have been able to

locate is that made by Johann Wahlberg in 1841 in KwaZulu-Natal Province (previously

Natal), South Africa and recorded as occurring between ‘Port Natal and Pieter. Mauritzburg’

(Wahlberg s.n.). The species was next collected by John Sanderson on 10 September 1858 in

‘Port Natal, Clairmont flat’ (Sanderson 49) and again in 1860 without a precise locality in

‘Natal’ (Sanderson 416). In October 1883, John Medley Wood made a collection in Clairmont

(Wood 1096). Based on the specimens collected in 1860 (Sanderson 416) and 1883 (Wood 1096),

Baker (1885) named and described the species. Wood made a further collection in 1892, again

in Clairmont (Wood 4662). In August of the following year, Friedrich Schlechter collected the

species near Clairmont, spelt as ‘Claremont’ on his label (Schlechter 3160). Using Schlechter’s

specimen, Baker (1904) described the new species Kniphofia pedicellata Baker. Berger (1908),

subsequently reduced K. pedicellata to a synonym of K. pauciflora.

The two collections (Sanderson 416 and Wood 1096) cited by Baker (1885, 1896) were treated by

Codd (1968, 2005) as syntypes. The Wahlberg s.n. and Sanderson 49 specimens were overlooked

by Baker (1885, 1896) and Berger (1908). The Wahlberg s.n. specimen was determined by Codd

in August 1961 and included in the citation of specimens in his treatment of Kniphofia Moench

(Codd 1968).

As part of our current taxonomic research on Kniphofia species in southern Africa that are under

threat and requiring conservation, we traced earlier known localities for species from herbarium

records. For K. pauciflora, minor problems amongst the syntypes and duplicates were noted and

it was found to be the only member of the genus not previously typified. We found it necessary

to lectotypify the species to specify a single specimen and to clarify the status of the duplicates.

In this note, we also provide an account of the last remaining population of K. pauciflora at the

Clairwood Racecourse, Durban, South Africa.

Research method and design

A thorough review of the literature pertaining to this species was conducted (see ‘Introduction’

as well as ‘Results and discussion’). Herbarium material, either actual specimens or images, were

examined for the type and include collections housed in the Natural History Museum, London

(BM), Bolus Herbarium, University of Cape Town, Cape Town (BOL), Royal Botanic Garden

Edinburgh, Edinburgh (E), Royal Botanic Gardens, Kew (K), KwaZulu-Natal Herbarium, South

African National Biodiversity Institute, Durban (NH), National Herbarium, South African

National Biodiversity Institute, Pretoria (PRE), Swedish Museum of Natural History, Stockholm

(S), South African Museum Herbarium, South African National Biodiversity Institute, Cape

Town (SAM), Trinity College, Dublin (TCD) and Universitat Zurich, Ztirich (Z) (herbarium

acronyms follow Holmgren, Holmgren and Barnett [1990)).

doi:10.4102/abc.v44i1.151

aaa oe Page 2 of 4 Eo Short Note -~

Results and discussion

Lectotypification of Kniphofia pauciflora

The specimen Sanderson 416, at K, has two intact open flowers

and an undehisced capsule, whilst the duplicate at TCD

has two intact old flowers. This distorts the characteristic

inflorescence in K. pauciflora and does not match the

description. Furthermore, the Sanderson specimen lacks a

precise locality; information that is useful for determining

Kniphofia species in KwaZulu-Natal. Wood 1096 (K) is the

most appropriate specimen for a type of the species as it

best matches the description. There are two specimens on

the sheet with full inflorescences clearly representative of

the species. Specimens with the same collector number are

also held at NH and BOL but show discrepancies in the

collecting dates on their labels. The Kew Herbarium received

the specimen from John Medley Wood in April 1881 and this

is written on the specimen sheet as well as in the record of

plant determinations sent from the Director of K to NH. In the

Herbarium Catalogue (Numbers 1—1944) at NH (Figure 1a),

Wood 1096 is recorded as collected in October with no year

and is amongst entries collected during 1880, starting with

Wood number 790 and ending with 1187 for that year. The

label on the NH specimen in Wood’s handwriting indicates

the collection date as October 1883 (Figure 1b). The collection

date on the BOL specimen label is November 1886, again in

Wood’s handwriting (Figure 1c). It is inferred that Wood 1096

was collected in 1880 and received by K in 1881. Presumably

the specimens collected in October 1883 (NH) and November

1886 (BOL) were collected during return visits to the same

population and therefore allocated the collector number of the

year 1880.

The available evidence suggests that of the three specimens

numbered Wood 1096, only the K specimen collected in

1880 constitutes original material and is therefore available

for lectotypification. The NH and BOL collections must be

regarded as topotypes, without nomenclatural standing.

Kniphofia pauciflora Baker in Journal of Botany, London

23: 280 (1885); Baker: 65, t.10 (1892a); Baker: t.7269 (1892b);

Baker: 279 (1896); Mallett: 101, t.43 (1906); Berger: 41 (1908);

Codd: 438 (1968); Codd: t.1995 (1989); Codd: 31 (2005). Type:

South Africa, KwaZulu-Natal: ‘Natal, Claremont flat’, Wood

1096 (K—image!, lecto. designated here) (Figure 2, barcode/

id K000256208).

K. pedicellata Baker: 998 (1904). Type: South Africa, KwaZulu-

Natal: ‘Clairmont’, Schlechter 3160 (Z—image!, holo.).

Distribution and habitat

Kniphofia pauciflora is a South African endemic, restricted

to KwaZulu-Natal Province. It is currently known from a

single locality in Durban, namely the inner field of the track

in the Clairwood Racecourse. Prior to 1956, K. pauciflora was

known to have a wider distribution within the Durban area

but populations were reduced through urban development.

Codd (1968) makes reference to J.W. Reyburn’s observation

http://www.abcjournal.org/ 39 | doi:10.4102/abce.v44i1.151

of the Pinetown (Mariannhill) area as being drained and the

population could no longer be found.

Ecology

The species grows at 24 m.a.s.l. in seasonally waterlogged

grassland, in full sun (Figure 3a). The soil is dark-coloured

and sandy.

Conservation status

The status of K. pauciflora is ‘Critically Endangered’

according to Raimondo et al. (2009). The species was

b mare penitnnsen.

Merbariseay

Source: Reproduced with permission from Bolus Herbarium (BOL) and South African National

Biodiversity Institute, KwaZulu-Natal Herbarium (NH)

FIGURE 1: Kniphofia data depicting, (a) photograph of KwaZulu-Natal Herbarium

Catalogue showing entry for 1096 in Wood’s handwriting, (b) label dataon KwaZulu-

Natal Herbarium specimen and (c) label data on Bolus Herbarium specimen.

previously thought to be extinct (Hilton-Taylor 1996; Scott-

Shaw 1999). In September 2012, 21 clumps of plants were

recorded at Clairwood Racecourse, with an average of six

inflorescences per clump (Figure 3b). The population was

successful in its habitat as a result of the interest shown

by Gold Circle, the former owners of the racecourse, and

conservation efforts by the lead author since 1968. The

present owners of the racecourse are Capital Property Fund.

The habitat (and species) is now threatened by a proposed

logistics and distribution park to support the development

of a dug-out port at the former Durban International Airport

(South Durban Basin). This confirms its status as ‘Critically

Endangered’.

Other material examined

SOUTH AFRICA, KwaZulu-Natal: Port Natal [Durban],

Claremont flat, 10 Sept. 1858, Sanderson 49 (TCD); Clairmont,

Oct. 1880, Wood 1096 (K); Clairmont flat, Oct. 1883, Wood

1096 (NH); on Clairmont flat, Nov. 1886, Wood 1096 (BOL);

Clairmont, near Durban, 22 Sept. 1892, Wood 4662 (BOL, E,

NH, SAM, Z); near Claremont, Aug. 1893, Schlechter 3160

(Z, two sheets); Clairmont, 12 Sept. 1898, Wood 7293 (PRE);

Clairmont, 30 Sept. 1899, Wood 7639 (BM); Clairmont, 25

Sept. 1907, Wood 10495 (NH, Z); Clairwood Racecourse,

07 Sept. 1968, Baijnath 593 (UDW); Clairwood Racecourse,

ex. cult. May 1985, Baijnath s.n. PRE58798 (PRE); near

- Page 3 of 4 e Shoyt| NOt a =

Source: Reproduced with the consent of the Board of Trustees, Royal Botanic Gardens, Kew

FIGURE 2: Image of lectotype of Kniphofia pauciflora (Wood 1096).

Durban, Jan. 1914, Indian Collector s.n. NH15971/2 (NH,

two sheets); Port Natal and Pieter. Mauritzburg [Durban

and Pietermaritzburg], 1841, Wahlberg s.n. (S); Pinetown

District, Mariannhill, Oct. 1951, Reyburn s.n. NH40464 (NH);

Pinetown District, Mariannhill, Nov. 1954, Reyburn s.n.

NH42151, PRE 37002 (NH, PRE); Pinetown, ex cult., 12 Nov.

1956, Reyburn s.n. PRE 37003 (PRE); without precise locality,

1860, Sanderson 416 (K, TCD).

Acknowledgements

The Curators of BM, BOL, E, K, NH, PRE,S, SAM, TCD and Z

are thanked for providing images of K. pauciflora specimens.

Source: Photographs by H. Baijnath

FIGURE 3: Kniphofia pauciflora as seen growing within the inner field of the track

in the Clairwood Racecourse, Durban, South Africa, (a) habitat and (b) clumps.

http://www.abcjournal.org/ 40 fo hogy MOA COPA ep ope Pe va AU NS he pa =

Page 4 of 4

Dr Yashica Singh kindly allowed access to J.M. Wood’s

collecting records and communications with K held at NH.

South African National Biodiversity Institute’s Biodiversity

Information Management Directorate, Cape Town provided

PRECIS data and its Mary Gunn Library, Pretoria made

available earlier literature. University of KwaZulu-Natal is

thanked for research support. We are grateful to Gold Circle

staff for access to the Clairwood Racecourse site.

Competing interests

The authors declare that they have no financial or personal

relationships that may have inappropriately influenced

them in writing this article.

Authors’ contributions

H.B. (University of KwaZulu-Natal) was responsible for data

collection. Both H.B. and S.R. (University of KwaZulu-Natal)

contributed to the manuscript preparation.

References

Baker, J.G., 1885, ‘A synopsis of the Cape species of Kniphofia’, Journal of Botany

London 23, 275-281.

Baker, J.G., 1892a, ‘Kniphofia pauciflora’, Gardeners’ Chronicle ser. 3(12), 65, t. 10.

Baker, J.G., 1892b, ‘Kniphofia pauciflora’, Curtis’s Botanical Magazine ser. 3, 38, t.7269.

Baker, J.G., 1896, ‘Kniphofia’, in W.T. Thistleton-Dyer (ed.), Flora Capensis 6, pp.

275-285, Reeve, London.

Baker, J.G., 1904, ‘Liliaceae’, Bulletin de I’Herbier Boissier ser. 2(4), 996-1002.

Berger, A.W., 1908, ‘Liliacaeae—Asphodeloideae—Aloineae’, in A. Engler (ed.), Das

Pflanzenreich: Regni Vegetabilis Conspectus 38, Verlag von Wilhelm Engelmann,

Leipzig.

Codd, L.E., 1968, ‘The South African species of Kniphofia (Liliaceae)’, Bothalia 9,

363-511.

Codd, L.E., 1989, ‘Kniphofia pauciflora’, The Flowering Plants of Africa 50, 1995.

Codd, L.E., 2005, ‘Asphodelaceae (First Part): Kniphofia’, in G. Germishuizen & B.A.

Momberg (eds.), Flora of Southern Africa 5(1) fascicle 2, pp. 1-94, South African

National Biodiversity Institute, Pretoria.

Hilton-Taylor, C., 1996, ‘Red data list of southern African plants’, Strelitzia 4, South

African National Botanical Institute, Pretoria.

Holmgren, P.K., Holmgren, N.H. & Barnett, L.C., 1990, Index Herbariorum Part 1: The

herbaria of the world, New York Botanical Garden, New York.

Mallett, G.B., 1906, ‘Kniphofias and their culture’, Gardeners’ Chronicle ser. 3, 39, p. 101.

Raimondo, D., Von Staden, L., Foden, W., Victor, J.E., Helme, N.A., Turner, R.C. et

al., 2009, ‘Red list of South African plants’, Strelitzia 25, South African National

Biodiversity Institute, Pretoria.

Scott-Shaw, C.R., 1999, Rare and threatened plants of KwaZulu-Natal and

neighbouring regions, KwaZulu-Natal Nature Conservation — Service,

Pietermaritzburg.

http://www.abcjournal.org/

doi:10.4102/abc.v44i1.151

— oe Page 1 of a

Short Note ——

Nomenclatural adjustments in African plants 1

Authors:

Peter Goldblatt??

John C. Manning?

Affiliations:

‘Research Centre for Plant

Growth and Development,

University of KwaZulu-Natal,

South Africa

*B.A. Krukoff Curator

of African Botany,

Missouri Botanical Garden,

United States

>Compton Herbarium, South

African National Biodiversity

Institute, South Africa

Correspondence to:

John Manning

Email:

j-manning@sanbi.org.za

Postal address:

Private Bag X7, Claremont

7735, South Africa

Dates:

Published: 10 Dec. 2014

How to cite this article:

Goldblatt, P. & Manning,

J.C., 2014, ‘Nomenclatural

adjustments in African

plants 1’, Bothalia 44(1), Art.

#169, 9 pages. http://dx.doi.

org/10.4102/abc.v44i1.169

Licensee: AOSIS

OpenJournals. This work is

licensed under the Creative

Commons Attribution License.

Read online:

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Background: Ongoing systematic studies of the African flora necessitate periodic nomenclatural

adjustments and corrections.

Objectives: To effect requisite nomenclatural changes.

Method: Relevant literature was surveyed and type material located and studied.

Results: Nomenclatural corrections were published in Justicia L. (Acanthaceae), Babiana Ker

Gawl. and Geissorhiza Ker Gawl. (Iridaceae) and Zaluzianskya F.W.Schmidt (Scrophulariaceae).

Conclusions: Firstly, a complete enumeration of all southern African species of Justicia was

provided within the infrageneric classification for the genus accepted by Graham (1988) and

later modified and expanded by Ensermu (1990) and Hedrén (1990). In this circumscription,

Justicia includes such well-established segregates in an African context as Adhatoda Miller,

Aulojusticia Lindau, Duvernoia E.Mey. ex Nees, Monechma Hochst. and Siphonoglossa Oersted.

Both southern African species of Adhatoda were transferred to Justicia, as well as all of the

southern African species of Monechma, with eight new combinations or replacement names

provided. All species were placed to section within Justicia. Secondly, the type of Gladiolus

nervosus Lam. (1788) was considered to be conspecific with Gladiolus strictus Aiton (1789)

and is therefore the earliest available name for the species currently known as Babiana stricta

(Aiton) Ker Gawl. The new combination Babiana nervosa (Lam.) Goldblatt & J.C-Manning was

provided. Thirdly, Geissorhiza ornithogaloides has been regarded as a new species described

by EW. Klatt (1866) but the name should be treated as the combination G. ornithogaloides

(Lichst. ex Roem. & Schult.) Klatt, based on Ixia ornithogaloides Lichst. ex Roem. & Schult.

(1817a). Examination of the type showed that it is conspecific with Geissorhiza marlothi

R.C.Foster (1941) and it is therefore the valid name for the taxon treated as G. ornithogaloides

subsp. marlothit (R.C.Foster) Goldblatt. An epitype for the taxon was designated and the new

combination G. ornithogaloides subsp. flava (Klatt) Goldblatt & J.C.Manning was provided for

the taxon currently treated as G. ornithogaloides subsp. ornithogaloides. Lastly, Reyemia Hilliard

was reduced to a synonym of Zaluziaskya as sect. Reyemia (Hilliard) J-C.Manning & Goldblatt

and the new combination Zaluziaskya chasmanthiflora (Hilliard) J.C.Manning & Goldblatt was

provided. A neotype was selected for Zaluziaskya nemesioides Diels.

1. Synopsis of the genus Justicia L. (Acanthaceae) in

southern Africa, with eight new combinations for species

of Adhatoda Miller and Monechma Hochst.

Authors: John C. Manning and Peter Goldblatt

Dates:

Received: 02 Oct. 2013

Accepted: 14 Nov. 2013

Introduction

Justicia L. (Acanthaceae: Ruellieae: Justiciinae) (Scotland & Vollesen 2000) is the largest genus of

Acanthaceae, containing at a conservative estimate some 600 spp. (Graham 1988). As noted by

Graham (1988) in her landmark review of the delimitation of Justicia, the circumscription of the

genus has historically followed one of two divergent trends — either the adoption of a broad view

of the genus, or towards the recognition of greater or lesser numbers of small segregate genera.

The worldwide distribution of Justicia and the high degree of morphological variation amongst

the species has hampered the general acceptance of one or the other of these two options, with

workers generally focused on regional floristic accounts and thus missing the taxonomic wood for

the trees. The result has been a confusing situation, whereby some regional floras recognise one or

more of the segregate genera whilst others do not. The need for a stable, or at least comprehensive,

10i:10.4102/abc.v44i1.169.§ ——_—

SSS

classification of the genus stimulated Graham (1988) to

undertake the first worldwide morphological survey of

Justicia since Lindau’s (1895) revision for Die natutirlichen

Pflanzenfamilien. The result of her extensive study of

vegetative and floral morphology and pollen ultrastruture

was the conviction that the genus is most appropriately and

satisfactorily circumscribed in the broad sense favoured by

Lindau (1895), thus including such well-established segregates

in an African context as Adhatoda Miller, Aulojusticia Lindau,

Duvernoia E.Mey. ex Nees and Siphonoglossa Oersted.

The resulting infrageneric classification for Justicia proposed

by Graham (1988) recognised sixteen sections, seven of them

in southern Africa, and enumerated 295 species, representing

an estimated half of the species in the genus. Amongst

the species not listed were several from southern Africa.

Although clearly not complete, Graham’s classification

forms the basis for future taxonomic studies in the genus

and a framework in which to incorporate additional species.

Monechma Hochst. was retained by Graham (1988) as distinct

from Justicia. The genus is traditionally diagnosed by its

two-seeded capsules containing smooth, compressed seeds;

whereas, Justicia is characterised by having mainly four-

seeded (rarely two-seeded) capsules containing at most

weakly compressed, mostly rugose (rarely smooth) seeds.

The validity of this distinction in the light of the occurrence

of intermediate forms was questioned by Hedrén (1990),

who concluded that the two taxa could not be upheld at

generic level. Accordingly, he reduced Monechma to a section

within Justicia, as had been done earlier by Lindau (1895),

thus increasing the number of sections in southern Africa

to eight. In the same year, J. subsect. Ansellia (C.B.Clarke)

V.A.W.Graham was raised by Ensermu (1990) back to

sectional level as sect. Ansellia C.B.Clarke.

Graham’s (1988) classification of Justicia was not implemented

for the floristic treatment of the tribe for southern A frica (Baden

etal. 1995) but has been adopted in later regional treatments for

sub-Saharan Africa (Scotland & Vollesen 2000). As a result, the

generic circumscriptions currently accepted for the southern

African subregion are now out of line with those for the Flora

of tropical East Africa (Darbyshire, Vollesen & Kelbessa 2010)

and the Flora of Somalia (Hedrén & Thulin 2006) to name just

two, both of which adopt the broad circumscription of Justicia

accepted by Graham (1988) and Hedrén (1990), as will the

planned volume for the Flora Zambesiaca region.

A start at integrating the treatment of the southern African

flora with recent accounts for the rest of the continent was

made by the formal transfer to Justicia of the remaining species

of Aulojusticia (= Siphonolossa spp.) by Manning and Goldblatt

(2012) and Duvernoia by Cubey (2006). What remains to be

effected is the transfer to Justicia of the two southern African

species of Adhatoda and all of the southern African species

of Monechma, several of which require new combinations

or replacement names, as well as the placement within the

appropriate sections in Graham’s (1988) classification of the

southern African species that were left unplaced by her.

- http://www.abcjournal.org 43 | doi:10.4102/abc.v44i1.169

We do this here and provide a complete enumeration of the

southern African species within the infrageneric classification

developed by Graham (1988) and modified and expanded

by Ensermu (1990) and Hedrén (1990). This will facilitate the

uniform curation of the southern African material in a system

compatible with that adopted elsewhere in sub-Saharan

Africa.

Results

The primary characters used by Graham (1988) to circumscribe

her sections are the structure of the inflorescence and the

pollen, notably the number of apertures and the condition

of the margocolpi. Placing the currently unplaced species

in an appropriate section in the system was mostly

straightforward through direct association with allied

species already in the system. Notable exceptions to

this are the two southern African species of Adhatoda,

A. andromeda (Lindau) C.B.Clarke and A. densiflora (Hochst.)

J.C.Manning. Their association with the genus Adhatoda

dates from the treatment of the family for Flora capensis by

Clarke (1901), who included them in Adahatoda rather than

in Justicia on account of their pollen having entire rather

than areolate margocolpi, the latter then considered to be

diagnostic for Justicia. Their placement in Adhatoda was

later questioned by Manning and Getliffe-Norris (1995),

who considered that they merited segregation from the

tropical African and Indian members of the genus. These

species, including the type of Adhatoda, were assigned by

Graham (1988) to sect. Vasica, diagnosed by a shrubby

habit and 2-aperturate pollen.

The South African Adhatoda species, both perennial herbs

with 3-aperturate pollen, would certainly be misplaced in

sect. Vasica and conform best to the small sect. Rhaphidospora

(Nees) T.Anderson, based on the combination of their

contracted, cymose infloresences and 3-aperturate pollen

with entire margocolpi. This pollen type is otherwise found

only in sect. Betonica and in the New World sect. Drejerella,

both of which have strictly simple, spike-like inflorescences.

The pollen type in Adhatoda andromeda and Adhatoda densiflora

is evidently ancestral in Justicia (Graham 1988) and thus not

phylogenetically informative, whilst the highly condensed

inflorescences are difficult to interpret. Itis possible that the two

species are better segregated in a separate section but a trend

towards a similar contraction of the inflorescence is evident

in Justicia bolusit C.B.Clarke, which seems correctly placed in

this section, and we include them here pending evidence to the

contrary.

We place Justicia minima A.Meeuse, unusual amongst African

species in its 4-fid calyx, in sect. Tyloglossa, largely on its

similarity in inflorescence and pollen morphology to Justicia

linearispica, placed here by Graham (1988).

Our placement in sect. Harniaria of the species previously

treated as the genus Aulojustcia, as recommended by

Hedrén (1990), is entirely consistent with the derived,

sessile inflorescences and 2-aperturate pollen with areolate

margocolpi that define this section. Similarly, placement of

Justicia cuneata Vahl. in the same section accords with its

sessile, 1-flowered axillary cymes and areolate margocolpi.

The southern African species of Monechma as treated by

Immelman (1995) were placed en bloc in sect. Monechma,

following Hedrén (1990). Inflorescence morphology in the

section is very variable (Hedrén 1990), raising the possibility

that the section is not monophyletic, although all species

examined share derived 2-aperturate pollen with areolate

margocolpi.

Taxonomic treatment

Justicia L., Species plantarum: 15 (1753).

Type: J. hyssopifolia L., lecto., designated by Hitchcock &

Green (1929), fide Graham (1988).

Description

Herbaceous or shrubby perennials or annuals. Leaves: opposite,

simple, entire. Inflorescence: a simple false-spike or compound

with dichasial or spike-like subunits. Flowers: each subtended

by a bract and two bracteoles (or bracts absent in simple

dichasia), white, yellow, pink to purple or red. Calyx: mostly

5-partite to near base, rarely + halfway, segments equal or

with 1 segment reduced or absent and then 4-partite. Corolla:

funnel-shaped or trumpet-shaped, bilabiate, upper lip usually

bifid (rarely entire), lower lip trilobed, throat usually pubescent

within. Stamens: 2, dithecous [rarely monothecous fide Graham

(1988) but not in the southern African taxa], thecae equal or

unequal, usually superposed, often oblique, lower usually with

sterile tail-like appendage. Pollen: 2 or 3(4) porate, subprolate

to perprolate, surface finely reticulate, trema ornamented with

4 or more margcocolpi, these either entire and band-like or

discontinuous and forming areoli or peninsulae. Ovary: with

2 ovules per locule, pubescent or glabrous; style lying within

channel (rugula) along upper lip; stigma minutely 2-lobed.

Capsules: + clavate with sterile stipe, 4-seeded, or 2-seeded

or 3-seeded by abortion, valves remaining entire during

dehiscence or rarely septum and adjacent capsule walls rising

(placenta elastic). Seeds: supported by retinacula, spheroidal

to discoid, testa smooth or variously ornamented. + 600 spp.,

worldwide in tropics and subtropics.

BOX 1: Key to Justicia L. (Acanthaceae) sections in southern Africa.

I. Sect. Vasica Lindau. Type: J. adhatoda L., lecto., designated

by Graham: 584 (1988).

Adhatoda Miller. Type: Adhatoda ‘zeylesianum’ Miller =

J. adhatoda L.

Duvernoia E.Mey. ex Nees. Type: D. adhatodoides

E.Mey. ex Nees = J. adhatodoides (E.Mey. ex Nees)

V.A.W.Graham.

Shrubby perennials. Inflorescence: a simple false-spike with

1 or 3 flowers per axil. Calyx: 5-partite with segments equal.

Corolla: 12 mm — 38 mm long, white or cream. Anther: thecae

equal or unequal, superposed and oblique, appendages

mostly small. Pollen: 2-aperturate, margocalpi areolate or

entire. Ovary: glabrous or pubescent. Capsules: sterile for

0.4-0.6 length. Seeds: scarcely or distinctly compressed,

rugulose-tuberculate-alveolate.

J. aconitiflora (A.Meeuse) Cubey [Duvernoia aconitiflora

A.Meeuse]

J. adhatodoides (E.Mey. ex Nees) V.A.W.Graham [Duvernoia

adhatodoides E.Mey. ex Nees, Adhatoda duvernoia C.B.Clarke,

nom illegit. superfl.]

II. Sect. Betonica (Nees) T.Anderson [Adhatoda sect. Betonica

Nees]. Type: J. betonica L., lecto., designated by Graham: 586

(1988).

Herbaceous or shrubby perennials. Inflorescence: spiciform,

1-sided with one flower per node, bracts conspicuous and

usually exceeding calyx, often + scarious. Calyx: 5-partite

with segments equal or 4-partite plus one reduced segment.

Corolla: 2.4 mm — 13.0 mm long, white, pale pink or blue.

Anther: thecae equal or slightly unequal, superposed

and oblique, appendages large or rarely lacking. Pollen:

3-aperturate, margocolpi entire. Ovary: glabrous or pubescent.

Capsules: sterile for + 0.4 length. Seeds: scarcely or distinctly

compressed, rugulose-tuberculate or appressed-pubescent.

J. betonica L.

J. betonicoides Burkill & C.B.Clarke

]. cheiranthifolia Nees

J. montis-salinarum A.Meeuse

2A. Mature capsules with one smooth, shiny seed per locule, always dehiscent;

1B. Inflorescences distinctly pedunculate or more than 7-flowered:

areolate

6B. Inflorescence spike-like; ovary glabrous:

pollen 2-aperturate, margocolpi areOlate ..........ccceccescssecceesccessseveseesssenserssessesensseens

2B. Mature capsules with two seeds per locule, rarely only one and then seeds either rough or capsules indehiscent; pollen various:

3As Inflorescences\sessile;polleni2-apentUratey mangocol pitajeolatewecsrscsseersesecessesereercreemesteretese cetera see ener tener meena ru es VI. sect. Harnieria

74. False-spike with 1—3(—15) flowers per node; pollen 3-aperturate, margocolpi areolate ...........cccceeeeceeseeseeseneeseesesees

78. False-spike with 1 flower per node; pollen 2-aperturate, margocolpi areolate

1A. Inflorescences sessile or very shortly pedunculate (peduncle < 10 mm long), 1-flowered to 6-flowered:

succes Veawousc ets sas aeeaauaeseaeduaevsuccvanissia taseaysePucusaatuvesd Wevevsevesneuasesecbvecen VII. sect. Monechma

3B. Inflorescences shortly pedunculate; pollen 2-aperturate, MargOCOlpi EMtire............sceccscescescssecceccscesencescsesesccesssecseuscuseasercasessuecssensessuseneeacucenecaeeee® V. sect. Justicia

4A. Inflorescence a simple false-spike (sometimes terminal and in adjacent axils giving a compound appearance); ovary pubescent; pollen margocolpi entire:

5A. Flowers two or more per node, the inflorescence thus cylindrical; bracts oblanceolate to elliptic, foliar; calyx lobes shorter than tube;

pollen 2-aperturate, MangOcolpi EMtine ...:...ccc.cnccevecsesecvectevscrevseesaccecessesserevseces

5B. Flowers mostly solitary at the nodes, the inflorescence thus one-sided; bracts ovate and + scarious with green veins; calyx lobes longer

than tube; pollen 3-aperturate, margocolpi ENtire............ccccecceeceeceseceeeeseeeeenee

4B. Inflorescence either compound and evidently cymose, or spike-like but then ovary glabrous; pollen margocolpi entire or areolate:

PES cea reece rece OS recone BLEEP Hack BE ARCCRee AERC en so -caceat ostr- OaeecE MoS soaked ass |. sect. Vasica

shack saset covaccatccuseencuce che ceataner th ceciee Seasnacte ines ti ean scons eUseeetencnor ste tateseereens Il. sect. Betonica

6A. Inflorescence evidently dichasial, the axis glandular; ovary mostly pubescent; pollen 3-aperturate, margocolpi entire or +

Ill. sect. Rhaphidospora

.. IV. sect. Tyloglossa

daddeta ct suvauaccectencvuacasWeccvdcopatcecansabevonctrast cueetase nents reren eb aereessacensktee Vill. sect. Ansellia

abcjournal.org

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Page 4 of 9

III. Sect. Rhaphidospora (Nees) T.Anderson [Rhaphidospora

Nees]. Type: J. glabra Koenig ex Roxb. = J. scandens Vahl

Shrubby or herbaceous perennials. Inflorescence: dichasial,

axis glandular. Calyx: 5-partite with segments equal. Corolla:

9 mm — 27 mm long, white or cream to pale pink. Anther:

thecae equal or slightly unequal, superposed and oblique,

appendages large. Pollen: 3-aperturate, margocalpi entire or

+ areolate. Ovary: mostly pubescent. Capsules: sterile for + 0.5

length. Seeds: scarcely or distinctly compressed, rugulose-

~ tuberculate or rarely echinate.

Group A: Shrubs; inflorescence a + open or contracted cyme.

J. bolusii C.B.Clarke

J. campylostemon (Nees) T.Anderson [Leptostachya campy-

lostemon Nees]

J. scandens Vahl

J. glabra Koenig ex Roxb.

Group B: Herbaceous perennials; inflorescence a highly

contracted, strobilate cyme. (Note: these two species

are placed here based on the combination of their highly

contracted dichasial inflorescences and 3-aperturate pollen

with entire margocolpi. This pollen type is otherwise found

only in sect. Befonica and in the New World sect. Drejerella,

both of which have strictly simple, spike-like inflorescences.)

J. andromeda (Lindau) J.C.Manning & Goldblatt, comb.

nov. Duvernoia andromeda Lindau in Engler & Prantl, Die

naturlichen Pflanzenfamilien 4 (3B): 339 (1895). Adhatoda

andromeda (Lindau) C.B.Clarke: 76 (1901).

J. densiflora (Hochst) J.C.Manning & Goldblatt, comb. nov.

Gendarussa densiflora Hochst. in Flora 28: 71 (1845). Adhatoda

densiflora (Hochst.) J.C.Manning in Manning & Getliffe-

Norris: 490 (1985).

IV. Sect. Tyloglossa (Hochst.) Lindau [Tyloglossa Hochst.].

Type: J. palustris (Hochst.) T.Anderson, lecto designated by

Graham: 590 (1988).

Herbaceous perennials. Inflorescence: compound but spike-like

with 1-3(-15) flower per node. Calyx: 5-partite with segments

equal (rarely 4-partite). Corolla: 5 mm — 12 mm long, white,

yellow or purple. Anther: thecae equal, superposed and

parallel or oblique, appendages large. Pollen: 3-aperturate,

margocolpi areolate. Ovary: glabrous. Capsules sterile for + 0.35

length. Seeds: scarcely compressed, ammonite-like or rugose.

J. flava (Vahl) Vahl [Dianthera flava Vahl]

J. fasciata Nees

J. kirkiana T.Anderson

J. minima A.Meeuse (Note: placed here on account of its

spike-like inflorescence and 3-aperturate pollen with areolate

margocolpi. Although the white flowers and 4-partite calyx

are anomalous in the section, L. linearispica C.B.Clarke from

tropical Africa, placed here by Graham (1988) as a ‘Peripheral

species’, has a very similar inflorescence and 4-partite calyx

with the fifth segment reduced.)

http://www.abcjournal.org 45 | doi:10.4102/abc.v44i1.169 ———

J. petiolaris (Nees) T.Anderson

subsp. petiolaris

subsp. bowiei (C.B.Clarke) Immelman [J. bowiei

C.B.Clarke]

subsp. incerta (C.B.Clarke) Immelman [J. incerta

C.B.Clarke]

V. Sect. Justicia

Shrubs or perennials. Inflorescence: a simple dichasium,

bracts absent. Calyx: 5-partite with segments equal. Corolla:

17mm -— 21 mm long, white or cream. Anther: thecae equal or

slightly unequal, level or slightly superposed and oblique,

appendages large. Pollen: 2-aperturate, margocolpi entire.

Ovary: glabrous. Capsules: sterile for + 0.5 length. Seeds: not

compressed, verrucose.

J. cordata (Nees) T.Anderson [Leptostachya cordata Nees]

J. guerkeana Schinz

J. orchioides L.f.

subsp. orchioides

subsp. glabrata Immelman

J. platysepala (S.Moore) P.G.Mey. [Monechma platysepala

S.Moore]

J. thymifolia (Nees) C.B.Clarke [Adhatoda thymifolia Nees]

VI. Sect. Harnieria (Solms-Laub.) Benth. [Harnieria Solms-

Laub.]. Type: H. dimorphocarpa Solms-Laub = J. heterocarpa

T.Anderson

Aulojusticia Lindau. Type: A. linifolia Lindau = J.

linifolia (Lindau) V.A.W.Graham

[Siphonoglossa pp. excl. type.]

Shrubs, herbaceous perennials or annuals. Inflorescence: a

sessile axillary cluster of + sessile flowers, bracts usually

absent. Calyx: 5-partite with segments equal. Corolla: funnel-

shaped or trumpet-shaped with short or elongate tube,

5 mm — 58 mm long, white or yellow to lilac or purple.

Anther: thecae equal, superposed and parallel or oblique,

appendages large. Pollen: 2-aperturate, margocolpi areolate.

Ovary: glabrous. Capsules: sterile for + 0.3 length. Seeds:

scarcely compressed, rugulose-tuberculate.

Group A: Corolla tube shorter than lips.

J. capensis Thunb.

J. cuneata Vahl (Note: placed here on account of its 1-flowered,

sessile axillary cymes and 2-aperturate pollen with areolate

margocolpi.)

subsp. cuneata

subsp. hoerleiniana (P.G.Mey.) Immelman _ [|].

hoerleiniana P.G.Mey.]

subsp. latifolia (Nees) Immelman [Gendarussa

orchoides subsp. latifolia Nees]

J. heterocarpa T.Anderson

subsp. heterocarpa

subsp. dinteri (S.Moore) Hedrén [J. dinteri S.Moore]

= = PERO SCS

J. odora (Forssk.) Vahl [Dianthera odora Forssk.]

J. protracta (Nees) T.Anderson [Gendarussa protracta Nees]

J]. kraussii C.B.Clarke

subsp. protracta

subsp. rhodesiana (S.Moore) Immelman [J. rhodesiana

S.Moore]

J. puberula Immelman

Group B: Corolla tube longer than lips.

J. linifolia (Lindau) V.A.W.Graham [Aulojusticia linifolia

Lindau, Siphonoglossa linifolia (Lindau) C.B.Clarke]

J. nkandlaensis (Immelman) J.C.Manning & Goldblatt

[Siphonoglossa nkandlaensis Immelman]

J. tubulosa (Nees) T.Anderson [Adhatoda tubulosa Nees]

Gendarussa leptantha Nees, Siphonoglossa leptantha

(Nees) Immelman

subsp. tubulosa

subsp. late-ovata (C.B.Clarke) J.C.Manning &

Goldblatt [J. pulegioides var. late-ovata C.B.Clarke]

VII. Sect. Monechma (Hochst.) T. Anderson [Monechma Hochst.].

Type: J. bracteata (Hochst.) Zarb.

Shrubs or annual or perennial herbs. Inflorescence: a terminal

spike, axillary clusters, or scattered. Calyx: 5-partite with

segments equal. Corolla: 5 mm — 19 mm long, white, red

or yellow. Anther: thecae equal or the lower slightly larger,

superposed, appendages large. Pollen: 2-aperturate,

margocolpi areolate. Ovary: glabrous or almost so. Capsule:

sterile for 0.3-0.5 length. Seeds: compressed, smooth,

sometimes fringed.

J. callothamnum (Munday) J.C.Manning & Goldblatt, comb.

nov. Monechma callothamnum Munday in South African

Journal of Botany 53: 140 (1987).

J. cleomoides S.Moore [Monechma cleomoides (S.Moore)

C.B.Clarke]

J. crassiuscula (P.G.Mey.) J.C.Manning & Goldblatt, comb.

nov. Monechma crassiusculum P.G.Mey. in Mitteilungen der

Botanischen Staatssammlung Miinchen 3: 604 (1960).

J. debilis (Forssk.) Vahl. [Monechma debile (Forssk.) Nees]

J. desertorum Engl. |Monechma desertorum (Engl.) C.B.Clarke]

J. distichotrichum Lindau [Monechma_ distichotrichum

(Lindau) P.G.Mey.]

J. divaricata Licht. ex Roem. & Schult. [Monechma divaricatum

(Licht. ex Roem. & Schult.) C.B.Clarke [as ‘(Nees) C.B.Clarke’]

[Note: the formal publication of Lichtenstein’s manuscript

name ‘Justicia divaricata’ by Roemer and Schultes (1817a) has

been overlooked until now, and the later Adhatoda divaricata

Nees (1847) has been erroneously accepted as the basionym

for the taxon.]

J. dregei J.C.Manning & Goldblatt, nom. nov. pro

Monechma mollissimum (Nees) P.G.Mey. in Mitteilungen

der Botanischen Staatssammlung Munchen 2: 304

(1957), non Justicia mollissima (Nees) Y.F.Deng &

T.F.Daniel (2011). Adhatoda mollisima Nees: 391 (1847).

(Note: the replacement epithet honours J.F. Drege who

collected the type.)

Dcjournal.or;

J. fleckii J.C.Manning & Goldblatt, nom. nov. pro Monechma

grandiflorum Schinz in Vierteljahrsschrift der Naturforschenden

Gesellschaft in Zurich 61: 441 (1916), non J. grandiflora Dum.

Cours. (1811). (Note: the replacement epithet honours E. Fleck

who collected the type.)

J. genistifolia Engl. [Monechma genistifolium (Engl.)

C.B.Clarke]

subsp. genistifolia

subsp. australe (P:G.Mey.) J-C.Manning & Goldblatt,

comb. nov. Monechma australe P.G.Mey. in Mitteilungen

der Botanischen Staatssammlung Miinchen 3: 602 (1960).

[Monechma genistifolium subsp. australe (P.G.Mey.) Munday]

J. incana (Nees) T.Anderson [Gendarussa incana Nees]

J. karroica J.C.Manning & Goldblatt, nom. nov. pro Monechma

robustum Bond in Journal of South African Botany 6: 67

(1940), non Justicia robusta T.Anderson ex C.B.Clarke (1900).

J. leucoderme Schinz [Monechma leucoderme (Schinz.)

C.B.Clarke]

J. namibensis J.C.Manning & Goldblatt, nom. nov. pro

Monechma calcaratum Schinz in Vierteljahrsschrift der

Naturforschenden Gesellschaft in Zurich 61: 441 (1916), non

J. calcarata Wall. (1830), nee Hochst. (1843).

J. salsola S.Moore [Monechma salsola (S.Moore) C.B.Clarke]

J. saxatilis (Munday) J.C.Manning & Goldblatt, comb. nov.

Monechma saxatile Munday in South African Journal of

Botany 3: 363 (1984).

J. serotina (P.G.Mey.) J.C.Manning & Goldblatt, comb. nov.

Monechma serotinum P.G.Mey. in Mitteilungen der Botanischen

Staatssammlung Munchen 11: 112 (1973).

J. spartioides T.Anderson [Monechma spartioides (T.Anderson)

C.B.Clarke]

J. tonsum (P.G.Mey.) J.C.Manning & Goldblatt, comb. nov.

Monechma tonsum P.G.Mey. in Mitteilungen der Botanischen

Staatssammlung Munchen 2: 304 (1957).

VII. Sect. Anselliana C.B.Clarke. [Justicia sect. Rostellaria

subsect. Anselliana (C.B.Clarke) V.A.W.Graham]. Type:

J. anselliana (Nees) T.Anderson, lecto., designated by Graham:

598 (1988).

Herbaceous perennials or rarely shrubs. Inflorescence:

a simple false-spike with 1 flower per node. Calyx:

5-partite with segments equal. Corolla: 5 mm — 7 mm

long, white or lilac. Anther: thecae equal, superposed and

parallel or oblique, appendages large. Pollen: 2-aperturate,

margocolpi areolate. Ovary: glabrous. Capsule: sterile

for + 0.3 length. Seeds: scarcely compressed, rugulose or

reticulate-alveolate.

J. anagalloides (Nees) T.Anderson [Adhatoda anagalloides Nees]

J. anselliana (Nees) T.Anderson [Adhatoda anselliana Nees]

J. exigua S.Moore

J. matammensis (Schweinf.) Oliv. [Adhatoda matammensis

Schweinf.]

Acknowledgements

We are most grateful to the referees for their valuable

comments.

; 46 doi:10.4102/abc.v44i1.169 ~

ARR

Competing interests

The authors declare that they have no financial or personal

relationships that may have inappropriately influenced them

in writing this article.

Authors’ contributions

Both authors collaborated on all aspects of the research.

2. Nomenclatural corrections

in Babiana and Geissorhiza

(Iridaceae: Crocoideae)

Authors: Peter Goldblatt and John C. Manning

Dates:

Received: 02 Oct. 2013

Accepted: 20 Jan. 2014

Introduction

Babiana Ker Gawl. (93 spp.) (Goldblatt & Manning 2007,

2012) is relatively widespread in southern Africa and

centred in the winter rainfall zone in the west of the

subcontinent. One of the better known species in Western

Cape Province, Babiana stricta (Aiton) Ker Gawl. (1803), is

a combination based on Gladiolus strictus Aiton (1789). The

location of a type, if it exists, is unknown and Goldblatt

and Manning (2004) designated the illustration in Curtis’s

Botanical Magazine accompanying the combination as a

neotype. Lewis (1959), in her monograph of Babiana, cited

Gladiolus nervosus Lam. (1788) as a synonym of B. stricta and

used the later epithet stricta for the species. Her apparently

inexplicable action is perhaps due to the presence of

the name Babiana nervosa Ker Gawl. in Index Kewensis,

which we assume she would have seen. Checking the

reference therein, we find neither G. nervosus nor B. nervosa

mentioned. In our accounts of Babiana we overlooked

Lewis’s error and maintained B. stricta, although in

somewhat revised circumscription (Goldblatt & Manning

2004, 2007). The type of G. nervosus, cultivated at the Jardin

Botanique in Paris, was grown from material collected by

Pierre Sonnerat at the Cape, which he visited briefly en

route to and from Mauritius in 1773 and 1781 (Gunn &

Codd 1981). The specimen is well preserved and is without

doubt the species known as B. stricta, as evinced by the stiff,

acute, upright leaves of that species, flowers with a narrow,

straight perianth tube + as long as the tepals, inner floral

bracts divided to the base, a pubescent ovary and most

significant, arrow-shaped anthers wider at the base.

We note that Lewis’s circumscription of B. stricta included

four varieties: of these var. erectifolia (G.J.Lewis) G.J.Lewis

is included in B. nervosa, var. grandiflora G.J.Lewis is

now Babiana tubaeformis Goldblatt & J.C.Manning, var.

regia G.J.Lewis is Babiana regia (G.J.Lewis) Goldblatt &

J.C.Manning and var. sulphurea (Jacq.) Baker has been

excluded as the type cannot be associated with any known

species (Goldblatt & Manning 2007; Goldblatt, Manning &

Gereau 2008).

http://www.abcjournal.org doi:10.4102/abce.v44i1.169

Short Note ----r

Lamarck’s epithet nervosus is valid in Gladiolus and is available

in Babiana [indexes of plant names show the name B. nervosa

Ker Gawl. (1804) but it does not appear in the purported place

of publication nor in Ker Gawler’s earlier article dealing with

B. stricta]. We provide the new combination here and reduce

B. stricta to synonymy. For completeness we include the full

synonymy of the species.

Taxonomic treatment

Babiana nervosa (Lam.) Goldblatt & J.C.Manning, comb. nov.

Gladiolus nervosus Lam., Encyclopédie méthodique 2: 724

(1788). Type: South Africa, without precise locality, ex hort.

Paris, Sonnerat s.n. (P: Herb. Lamarck, holo.—digital image!).

Gladiolus strictus [Sol. in] Aiton: 63 (1789) syn. nov.

Babiana stricta (Aiton) Ker GawlL.: t. 621 (1803). Type: South

Africa, without precise locality or collector, illustration in

Ker GawlL.-: t. 621 (1803) [neotype designated by Goldblatt &

Manning: 94 (2004)].

Babiana erectifolia G.J.Lewis: 3 (1938). Babiana stricta var.

erectifolia G.J.Lewis: 43 (1959). Type: South Africa, [Western

Cape], Brand Vlei, near Worcester, Sept. 1932, G.J. Lewis s.n. as

Nat. Bot. Gard 2686/32 (BOL, holo.!).

Gladiolus ringens Thunb.: 186 (1800), hom. illegit. non

Andrews (1798). Type: South Africa, without precise locality,

Thunberg s.n. (UPS: Herb. Thunberg, syn.).

?Babiana caesia Eckl.: 32 (1827). Type: South Africa,

without precise locality, ?Ecklon s.n. (location unknown,

not at S).

Babiana flavocaesia Eckl.: 32 (1827). Type: South Africa,

vicinity of Stellenbosch, ?Ecklon s.n. (S, holo.!).

In Geissorhiza Ker Gawl. (100 spp.) (Goldblatt & Manning

2009, 2013), the nomenclature of G. ornithogaloides and its

two subspecies need adjustment. The name Geissorhiza

ornithogaloides is currently treated as a new species

described by F.W. Klatt in 1866 (Goldblatt 1985). We note,

however, that Klatt cited as a syntype, Lichtenstein s.n. from

the ‘Koue Bokkeveld’. This specimen is the type of Ixia

ornithogaloides Lichst. ex Roem. & Schult. (1817a). Although

Klatt did not cite a basionym for his species, it should be

treated as a combination according to the International Code

of Botanical Nomenclature Art. 41.4 (especially examples

7,8 and 10) as it is based in part on the same type (Greuter

ef al. 2000).

The identity and type locality of the Lichtenstein collection

matches what is currently G. ornithogaloides subsp. marlothii,

based on Geissorhiza marlothii R.C.Foster (1941). As a result,

the name subsp. marlothii must be replaced by subsp.

ornithogaloides and this taxon, in turn, requires a new name in

place of its current circumscription as subsp. ornithogaloides.

We accordingly replace it with the name subsp. flava based

on Geissorhiza flava Klatt (1882). The choice of Ecklon & Zeyher

Irid. 225 (51.8) from Caledon as lectotype of G. ornithogaloides

Klatt by Goldblatt (1985) falls away. We also cite an epitype

for G. ornithogaloides because the type specimens lack corms,

important in distinguishing the two subspecies. The revised

nomenclature is as follows.

Geissorhiza ornithogaloides (Licht. ex Roem. & Schult.) Klatt

in Linnaea 34: 656 (1866). Ixia ornithogaloides Licht. ex Roem.

& Schult.: 376 (1817b). Trichonema ornithogaloides (Licht. ex

Roem. & Schult.) A.Dietr.: 583 (1833). Type: South Africa,

[Western Cape], Koue Bokkeveld [Cold Bokkeveld], possibly

Nov. 1803, Lichtenstein s.n. (B, holo.!).

subsp. ornithogaloides

Geissorhiza marlothit R.C.Foster: 66 (1941), syn. nov.

G. ornithogaloides subsp. marlothii (R.C.Foster) Goldblatt: 325

(1985). Type: South Africa, [Western Cape], Cold Bokkeveld,

Houdenbeck, 850 m, Marloth 10612 (B, holo.!; NBG!, PRE!,

iso.); [Western Cape], Schoongesig, Ceres, Hanekom 1222

(NBG, epi., designated here).

subsp. flava (Klatt) Goldblatt & J-C-Manning, comb. & stat.

nov. Geissorhiza flava Klatt: 392 (1882). G. ornithogaloides var.

flava (Klatt) R.C.Foster: 68 (1941). Type: South Africa, [Western

Cape], without precise locality, Breutel s.n. (B, holo.!).

[Waitzia flava Reichb. in Klatt: 392 (1882), cited in

synonymy]

[Geissorhiza romuleoides Eckl: 27 (1827), nom. nud.]

Acknowledgements

P.B. Phillipson (Missouri Botanical Garden) kindly confirmed

details of the type of Gladiolus nervosus in the Paris Herbarium

[P] and R.E. Gereau (Missouri Botanical Garden) provided

nomenclatural advice.

Competing interests

The authors declare that they have no financial or personal

relationships that may have inappropriately influenced them

in writing this article.

Authors’ contributions

Both authors collaborated on all aspects of the research.

3. Reyemia included in Zaluzianskya

(Scrophulariaceae: Limoselleae) with

the new combination Zaluzianskya

chasmanthiflora (Hilliard)

J.C.Manning & Goldblatt

Authors: John C. Manning and Peter Goldblatt

Dates:

Received: 10 June 2014

Accepted: 17 Sept. 2014

Introduction

The genus Reyemia Hilliard (1992) (Scrophulariaceae:

Limoselleae) was established for two species of annual

herbs from the Hantam and Roggeveld Plateau in Northern

Cape Province, South Africa, neither of them well known.

Reyemia nemestoides (Diels) Hilliard (previously Zalusianskya

nemesioides Diels) has been collected several times along

the length of the Hantam-Roggeveld escarpment between

Short Note —— —— SSS

Calvinia and Sutherland, but Reyemia chasmanthiflora Hilliard

is still known only from the type locality near Williston.

The relationships of Reyemia were presumed by Hilliard

(1994) to lie with Zalusianskya F.W.Schmidt, with which

it shares a derived, shortly toothed and plicate calyx and

similar, centrally depressed, cushion-shaped seeds. It was

distinguished from Zaluzianskya by what was described as a

‘loosely paniculate’ inflorescence, a resupinate corolla witha

patch of clavate hairs inside the throat on the posticous side,

and with two stamens and two staminodes. Examination of

the specimens confirms that the inflorescence comprises

essentially spicate units, sometimes with the lower flowers

shortly pedicellate. Species of Zalusiaskya have simple or

sparsely branched, spicate or racemose inflorescences, non-

resupinate flowers, sometimes with a ring of clavate hairs in

the mouth, and either four or two stamens, usually without

staminodes (Hilliard 1992). The differences between the

two genera are thus centred on the strongly branched

inflorescence and more zygomorphic flowers in Reyemia.

An examination of phylogenetic relationships in Manuleae

and Selagineae by Kornhall, Heidari and Bremer (2001),

based on the plastid gene regions ndhF and trnL, confirmed

the close relationship between Reyemia and Zaluzianskya. This

analysis, although including just four species of Zaluzianskya

plus R. chasmanthiflora, provided the first indication that

the latter was, in fact, nested in Zaluzianskya. A subsequent,

more intensively sampled study of relationships within

Zaluzianskya by Archibald, Mort and Wolff (2005) used a

combination of nuclear ITS and plastid rps16 and trnL-F gene

regions from a sampling of 23 species of Zaluzianskya, plus

the second species of Reyemia, R. nemesioides, to produce the

first broad phylogenetic analysis of the genus. This study

confirmed that Reyemuia is deeply nested within Zaluzianskya,

as sister to the clade comprising those members of section

Holomeria (Benth.) Hilliard & Burtt with four fertile stamens.

The placement is entirely consistent with the morphological

similarities between the two genera noted by Hilliard

(1994) and supplemented by additional states identified by

Archibald et al. (2005), viz. filaments decurrent to the base of

the corolla tube and a ligulate stigma.

These findings led Archibald et al. (2005) to conclude that the

differences in floral morphology between the two genera are

no greater than exists amongst other species of Zaluzianskya

and that Reyemia should be merged with Zaluzianskya.

The entire corolla lobes in the former species of Reyemia

and in members of Zaluzianskya sect. Holomeria supports

the close relationship between the two groups suggested

by the molecular analysis of Archibald ef al. (2005). There

are indications, however, that sect. Holomeria may not be

monophyletic as currently defined because Zaluzianskya

benthamiana was retrieved as a member of sect. Zaluzianskya

subsect. Zaluzianskya. This species (plus Zaluzianskya diandra

which was not included in the analysis) are unique in sect.

Holomeria in having just two stamens, raising the possibility

that the entire corolla lobes in Z. benthamiana and the other

@ doi:10.4102/abc.v44i1.169 ——— Steet = a

Page 8 of 9

members of sect. Holomeria are convergent. This possibility,

combined with the distinctive inflorescence and flowers of the

former members of Reyemia, favours the retention of the two

species as the separate section Reyemia within Zaluzianskya.

We implement this finding here in preparation for the

forthcoming regional flora of the Karoo region. We also select

an appropriate neotype from the same locality to replace the

holotype of Zaluzianskya nemestoides in the Berlin Herbarium

which has been lost.

Research method and design

We consulted the relevant literature and implemented the

necessary nomenclatural conclusions. Type material is cited

from the Berlin Herbarium (B) and Compton Herbarium, South

African National Biodiversity Institute, Cape Town, South

Africa (acronyms after Holmgren, Holmgren & Barnett 1990).

Taxonomic treatment

Zaluzianskya F.W.Schmidt, Neue und Seltene Pflanzen: 11

(1793), nom. cons., non Zaluzianskia Necker (= Marsilea L.).

Type species: Zaluzianskya villosa F.W.Schmidt.

Sect. Reyemia (Hilliard) J.C.Manning & Goldblatt, syn. et

stat. nov. Reyemia Hilliard in Edinburgh Journal of Botany 49:

297 (1992). Type species: Reyemia chasmanthiflora Hilliard =

Zaluzianskya chasmanthiflora (Hilliard) J.C.Manning & Goldblatt.

Zaluzianskya chasmanthiflora (Hilliard) J.C.Manning

& Goldblatt, comb. nov. Reyemia chasmanthiflora Hilliard

in Edinburgh Journal of Botany 49: 297 (1992). Type:

South Africa, [Northern Cape, Williston], Farm Annexe

Kransfontein 721, 02 Sept. 1986, Cloete & Haselau 172 (NBG

[as STE], holo.).

Zaluzianskya nemesioides Diels in Botanische Jahrbticher

fur Systematik, Pflanzengeschichte und Pflanzengeographie

23: 482 (1896). Reyemia nemesioides (Diels) Hilliard: 297 (1992).

Type: South Africa, [Northern Cape], Hantam Mountains,

Meyer s.n. (Bt, holo.). Neotype, selected here: South Africa,

Northern Cape, Calvinia, gravel flats below Hantamsberg, 15

Sept. 1996, Goldblatt & Manning 10525 (NBG, neo.; MO, iso.).

Acknowledgments

Competing interests

The authors declare that they have no financial or personal

relationships that may have inappropriately influenced them

in writing this article.

Authors’ contributions

Both authors collaborated on all aspects of the research.

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http://www.abcjournal.org 50 | doi:10.4102/abc.v44i1.169 —— — i

Short Communication

Page 1 of 3

Schoenefeldia transiens (Poaceae): Rare new record

from the Limpopo Province, South Africa

Authors:

Aluoneswi C. Mashau?

Albie R. Gotze?

Affiliations:

1South African National

Biodiversity Institute,

Pretoria, South Africa

7Environment Research

Consulting, Potchefstroom,

South Africa

Correspondence to:

Aluoneswi Mashau

Email:

c.mashau@sanbi.org.za

Postal address:

Private Bag X101, Pretoria

0001, South Africa

Dates:

Received: 19 Aug. 2014

Accepted: 09 Sept. 2014

Published: 08 Dec. 2014

How to cite this article:

Mashau, A.C. & Gotze,

A.R., 2014, ‘Schoenefeldia

transiens (Poaceae): Rare

new record from the

Limpopo Province, South

Africa’, Bothalia 44(1), Art.

#325, 3 pages. http://dx.doi.

org/10.4102/abc.v44i1.325

Licensee: AOSIS

OpenJournals. This work is

licensed under the Creative

Commons Attribution

License.

Read online:

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Background: Schoenefeldia is a genus of C, grasses, consisting of two species in Africa,

Madagascar and India. It is the only representative of the genus found in southern Africa,

where it was previously only known from a few collections in the southern part of the Kruger

National Park (Mpumalanga Province, South Africa), dating from the early 1980s.

Objectives: The objective of this study was to document a newly recorded population of

Schoenefeldia transiens in an area that is exploited for coal mining.

Method: A specimen of S. transiens was collected between Musina and Pontdrift, about 30 km

east of Mapungubwe National Park, in the Limpopo Province of South Africa. The specimen

was identified at the National Herbarium (Pretoria).

Results: This is not only a new distribution record for the quarter degree grid (QDS: 2229BA),

but is also the first record of this grass in the Limpopo Province. The population of S. transiens

has already been fragmented and partially destroyed because of mining activities and is under

serious threat of total destruction.

Conclusion: It is proposed that the population of S. transiens must be considered to be of

conservation significance, and the population should be made a high priority in the overall

environmental management programme of the mining company that owns the land.

Introduction

Schoenefeldia Kunth (1830:283) is a genus of C, grasses, consisting of two species in Africa,

Madagascar and India (Watson & Dallwitz 1994). Schoenefeldia transiens (Pilger) Chiovenda

(Chiovenda 1916:186; Pilger 1914: 418) is known from Ethiopia, Somalia, Sudan, Kenya,

Tanzania, Uganda, Mozambique, Zimbabwe and South Africa (Clayton et al. 2014). It is the

only representative of the genus found in southern Africa, where it was previously only

known from a few collections in the southern part of the Kruger National Park (Mpumalanga

Province, South Africa), dating from the early 1980s (see details under ‘Additional specimens

examined’ below). No other specimens collected between 1982 and the current collection

(2014) could be found.

A specimen of S. transiens was collected between Musina and Pontdrift, about 30 km east of

Mapungubwe National Park, in the Limpopo Province of South Africa. This is not only a new

distribution record for the quarter degree grid (QDS: 2229BA), but the first record of this grass in

the Limpopo Province. As the collection reported on here was made in an area that is exploited by

coal mines, the conservation of this species in the Limpopo Province is not secure.

Taxonomic treatment

Schoenefeldia transiens (Pilg.) Chiov. in Resultati scientifici della missione Stefanini-Paoli nella

Somalia ttaliana. Le collezione botaniche 1:186 (1916).

Basionym: Chloris transiensis Pilg., in Botanische Jahrbiicher fiir Systematik, Pflanzengeschichte und

Pflanzengeographie 51:418 (1914).

Type: TANZANIA. Pare District: KwaSengiwa-Majiyajuu, Uhlig 882 (B100002186, holotype, e!).

Description

Densely tufted perennial grass, 700 mm — 1200 mm high. Leaf blade 350 mm x 5 mm. Inflorescence

of 2-4 digitate racemes; racemes 130 mm — 200 mm long, with obviously secund spikelets, these

solitary at point of attachment to rachis. Spikelet 3.5 mm — 5.0 mm long, small in comparison to

awns; florets 1 or 2, fertile lemma awn 10 mm — 25 mm, sterile lemma awn 25 mm — 45 mm long,

a http://www.abcjournal.org 51 | dol O'4102/abev44i1.32 5)

Ne,

{

| ow ray

Wigan

Sara

Source: Sample from Ellis 3548 (PRE), photographed by A.C. Mashau

Scale bar: (b), 2 mm.

FIGURE 1: Schoenefeldia transiens, depicted in, (a) habit (the growth form of the

plant) and (b) lemma (the lower of two bracts enclosing the grass flower or floret).

awns flexuous, curving gracefully, becoming entangled with

awns of other spikelets (Cope 1999; Gibbs Russell ef al. 1990);

anther 0.7 mm — 1.0 mm long (Figure 1). Flowering January

to February. Reported to be cleistogamous.

Distribution and ecology

Schoenefeldia transiens occurs in southern Africa in the

Mpumalanga and Limpopo Provinces of South Africa, and

northwards through to East Africa, Ethiopia, Somalia and

Sudan (Figure 2). The newly recorded S. transiens population

in the Limpopo Province is situated in the Limpopo River

Source: Created by H.M. Steyn

FIGURE 2: Known distribution range (@) of Schoenefeldia transiens in southern

Africa, with the black arrow indicating that further populations can be found to

the north.

Valley between Musina and Pontdrift, on the farm Over Vlakte

125 MS. It grows on a north-facing hill slope approximately

1 km south of the Limpopo River. Coordinates of specimens

and sub-populations of the larger population of S. transiens on

the farm Over Vlakte 125 MS in the Limpopo River Valley are

22°08'58.3" S, 29°40’ 49.7’ E and 22°09'01.9"S, 29° 40'39.6" E.

The new population of S. transiens occurs in Colophospermum

mopane woodland on relatively seasonally flooded flats, in

heavy soil, such as shallow loamy sandy soil that is associated

with a high percentage of sandstone and quartzite surface

rocks and outcrops (Figure 3). The habitat slope is between 5°

and 10°, with a predominantly northern aspect. The woody

component of the habitat is dominated by trees and shrubs

that are mostly between 2.2 m and 0.8 m (average 1.5 m)

high. The grass and herbaceous layer is only moderately to

poorly developed due mainly to the shallow rocky nature of

the substrate.

Other perennial grasses of significant abundance that occur

together with S. transiens are Fingerhuthia africana, Digitaria

eriantha, Stipagrostis uniplumis and Sporobolus nitens. Together

with C. mopane, the woody vegetation is dominated by Gardenia

resiniflua subsp. resiniflua, Terminalia prunioides, Ximenia

americana, Combretum apiculatum and Croton gratissimus var.

subgratissimus. Dominant herbs include Barleria virgula, Seddera

suffruticosa, Decorsea schlechteri, Hibiscus micranthus, Xerophyta

humilis, Tephrosia polystachya and Indigofera nebrowniana.

Conservation status and habitat sensitivity

The frequency of S. fransiens in southern Africa is rare.

Although the habitat of the new population itself is not

considered to be particularly sensitive, it is under imminent

threat of destruction as a result of current open cast coal

mining activities in the direct vicinity of the newly recorded

- http://www.abcjournal.org 52 | doi:10.4102/abe.v44i1.325 — a s

Source: Image (a) photographed by G. de Beer; (b) by A.R. Gotze

FIGURE 3: Schoenefeldia transiens (a) tall grass in foreground, (b) occurring in the rocky Colophospermum mopane wooland.

population. A strong suspicion exists that the population has

already been fragmented and partially destroyed because

of mining activities. Depending on the future planning of

mining activities, the whole population is under serious

threat of total destruction. It is therefore proposed that this

population of S. transiens be considered to be of conservation

significance, and it should be made a high priority in the

overall environmental management programme of the mining

company that owns the land. It is the view of the authors

that if this population is lost as a result of carelessness or

ignorance by either the relevant conservation authorities or

the current landowner (i.e. the mining company), it will be a

loss not only to our natural heritage, but also a loss in terms

of the gene pool of this species and to the biodiversity of the

southern African region.

New collection record

SOUTH AFRICA. Limpopo: Limpopo River Valley, farm Over

Vlakte 125 MS (QDS: 2229BA), 24 April 2014, Gotzel 355 (PRE)).

Additional specimens examined

SOUTH AFRICA. Mpumalanga: Kruger National Park, 10 km

east of Satara Camp, Nwanedzi road at Msasame windmill

(QDS: 2431BD), 13 Jan. 1981, Ellis 3548 (PRE!); 12 km east of

Satara along Nwanedzi river road, 28 Jan. 1982, Ellis 3867

(PRE!); 6 km east of Satara along Nwanedzi river road, 31 Jan.

1981, Ellis 3542; 3543 (PRE!).

Acknowledgements

Our grateful thanks to: Hester Steyn, National Herbarium,

Pretoria, South African National Biodiversity Institute, for

http://www.abcjournal_org 53 | doi:10.4102/abc v44i1.325

the distribution map; Ronell Klopper, South African National

Biodiversity Institute, for comments on an earlier draft of

this manuscript; the South African National Biodiversity

Institute and Environment Research Consulting (ERC) for

support.

Competing interests

The author declares that she has no financial or personal

relationships that may have inappropriately influenced her

in writing this article.

Authors’ contributions

A.C.M. (South African National Biodiversity Institute/

University of Pretoria) prepared the draft and analysed the

data, A.R.G. (Environment Research Consulting) collected plant

material and made conceptual contributions.

References

Chiovenda, E., 1916, ‘Resultati scientifici della missione Stefanini-Paoli nella Somalia

italiana’, Le collezione botaniche 1, 186.

Clayton, W.D., Govaerts, R., Harman, K.T., Williamson, H. & Vorontsova, M., 2014,

World checklist of Poaceae, Royal Botanic Gardens, Kew, viewed 12 August 2014,

from http://apps.kew.org/wcsp/

Cope, T.A., 1999, ‘Gramineae’, in G.V. Pope (ed.), Flora zambesiaca 10(2), pp. 243-244,

Royal Botanic Gardens, Kew, London.

Gibbs Russell, G.E., Watson, L., Koekemoer, M., Smook, L., Barker, N.P., Anderson,

H.M. et a/., 1990, ‘Grasses of southern Africa’, Memoirs of the Botanical Survey

of South Africa 58, 291.

Kunth, C.S., 1830, Révision des Graminées 17, p. 282, t. 53.

Pilger, R.K.F., 1914, ‘Graminae africanae XII’, Botanische Jahrbucher fiir Systematik,

Pflanzengeschichte und Pflanzengeographie 51, 412-422.

Watson, L. & Dallwitz, M.J., 1994, The grass genera of the world, revised edn., CAB

International, Wallingford.

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