Plant endemism in Griqualand West, South Africa

A.W. Frisbya*, S.J. Siebertb, M. Struwigb, D.P. Cilliersb

aDepartment of Plant and Soil Sciences, University of Pretoria, Private Bag X20, Hatfield, Pretoria
0028, South Africa
bUnit for Environmental Sciences and Management, North-West University, Private Bag X6001,
Potchefstroom 2520, South Africa

*Corresponding author: Department of Plant and Soil Sciences, University of Pretoria, Private Bag X20,
Hatfield, Pretoria 0028, South Africa. Tel: 012 420 6031, Email: Arnold.Frisby@up.ac.za (A.W. Frisby)

Highlights

• Twenty-two species have their distribution ranges restricted to the Griqualand West region,

representing 1.4% of the region's flora.

• The presence of the hypothesized Griqualand West Centre of plant endemism is

confirmed.

• Many GWC endemics show indications of holo-endemism owing to their preference for the

Ca-rich substrates of the Ghaap Plateau.

• This study proposes the concept of ‘core area’ which highlights the area with the highest

endemic plant density.

Abstract
Griqualand West, a region in the semi-arid Northern Cape and North-West provinces of South
Africa, has been variously suggested to contain a number of range-restricted plant species,
and was proposed to be a local centre of plant endemism. The Griqualand West Centre
(GWC), hitherto demarcated by geological features and limited floristic data, is hereby
investigated to determine the true levels of endemism and its extent of occurrence. Findings
suggest that at least 23 plant species have their natural distribution ranges restricted to the
Griqualand West region. These endemics represent 1.4% of the region’s flora. Although this
is a lower than the predicted level of endemism, it matches the trends of endemicity found in
other centres in semi-arid savanna of southern Africa. Many of the GWC endemics show
indications of holo-endemism owing to their apparent preference for the Ca- and Mg-rich

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substrates of the Ghaap Plateau. When the total distributions of all GWC endemic species are
considered, then the resulting boundary of the GWC is more extensive than the substantial
area already proposed previously (>40 000 km2). This study therefore proposes the concept
of ‘core area’ in which distant outlier populations of endemic species (>100 km outside the
main distribution range with no suitable habitat in between) are discarded during the
demarcation of the centre’s boundary. It is proposed that this concept is best applied when
assessing extensive areas with few endemic species. A more refined GWC core area will allow
for more effective conservation and future research efforts by focussing attention on those
areas where high numbers of endemic plant species co-occur. Within the GWC core area,
specific regions, such as the increasingly densely populated Kimberley region, the banded
ironstone hill ranges, and the unique environment that is the Ghaap Plateau, are highlighted
as areas of conservation importance.

Keywords: centre of endemism; dolomite; endemic; Ghaap; ironstone; limestone; near-

endemic.

1. Introduction
Worldwide, many vascular land plants exhibit total distributions that are restricted to
small geographic regions (White, 1983; Van Wyk and Smith, 2001). In many cases, it has
been noted that many restricted-range (endemic) plant species share a common geographical
range (Jansson, 2003). Such regions are often referred to as local or regional “centres of plant
endemism” (Van Wyk and Smith, 2001). Centres of endemism are of interest for a wide
number of reasons, including, among others, directing conservation efforts (e.g. Anderson,
2002), understanding plant gene-flow and speciation (Ellstrand 1992; Anderson et al., 2001),
and assessing the potential impacts of climate change on endemic plant diversity (Loarie et
al., 2008; Dirnböck et al., 2011). In the diverse botanical region of southern Africa, various
centres of plant endemism have been identified or proposed (e.g. Van Wyk, 1996; Siebert et
al., 2002; Clark et al., 2009; Williamson and Balkwill, 2015; Hahn, 2017). Most of these are
associated with the Great Escarpment of southern Africa (Clark et al., 2011).
One region of semi-arid savanna in north-central South Africa not associated with the
escarpment, known as Griqualand West, has been noted as botanically diverse and rich in
endemic plants (Wilman, 1946; Acocks, 1988). Despite these reports, the patterns of floristic
diversity in Griqualand West are poorly known because of a sparsity of plant collections. Van
Wyk and Smith (2001) tentatively proposed that the region may be a local centre of plant
endemism, namely the Griqualand West Centre (GWC). This proposal was made in light of
the apparent presence of 40 range-restricted plant species in the region, representing
approximately 2.2% of the region’s flora (Van Wyk and Smith, 2001). Another factor that lead

2
to the proposal was Griqualand West’s diverse geology – a factor often known to be
associated with higher levels of plant endemism around the world (Kruckeberg and
Rabinowitz, 1985) and specifically in southern Africa (Wild, 1965; Siebert et al., 2001;
Williamson and Balkwill, 2015). However, as no phytogeographic study had been conducted
on the plants of Griqualand West, the possible presence of a centre of plant endemism in the
region has remained questionable. To address this shortcoming, this study attempted to
collate the known plant collections from the Griqualand West region to test the validity of the
proposed centre of endemism by analysing endemic plant distributions, the GWC’s proposed
borders and levels of floristic endemism. Such a study is crucial for the region from a
conservation perspective, as it is inadequately conserved, and under great pressure from
widespread mining activities which focus primarily on limestone, manganese ore and
ironstone deposits.

2. Materials and method

2.1 Study area
Griqualand West covers an area of around 40 000 km2, and is located in north-central
South Africa, primarily in the Northern Cape, but also extending into parts of North-West.
Griqualand West was inhabited by the Khoesaan from at least the Late Iron Age (Coon and
Hunt, 1965). The region is named after a Khoekhoe people inhabiting this area, namely the
Griqua (Van Wyk and Smith, 2001). The Griqua were semi-nomadic tribes with a diverse
heritage including Dutch, Tswana, Khoekoe and San people (Penn, 2005).
The topography of Griqualand West is made up of undulating landscapes with higher
and lower-lying areas. The eastern-central part of Griqualand West is an extensive flat plateau
approximately 130 km wide and 280 km long, known as the Ghaap Plateau (± 1 450 m a.s.l.).
It comprises primarily of dolomite and associated Ca-rich sediments. The western-central
section of the Griqualand West has a number of low mountain ranges and hills cutting through
the region from north to south, most prominent of which are the quarzitic Langberg (± 1 800
m a.s.l.) and the banded ironstone of the Kuruman (± 1 835 m a.s.l.) and Asbestos Hills (± 1
200 m a.s.l). The longest of these ranges is the Langberg stretching some 130 km (Raper et
al., 2014). Large intermontane valleys are filled with aeolian Kalahari sands.

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Table 1.
Selected climate variables illustrating the gradient from east to west and north to south in Griqualand
West.

Region Geographic Mean annual Mean annual Mean Mean

regions precipitation temperature annual frost annual soil
(mm) (°C) days (< moisture
0°C) stress (%)
Ghaap Plateau Eastern 411 17.1 42 83
Ironstone Hills Central 370 17.1 38 84
Langberg Western 292 17 35 84
Above combined Southern 358 17 38 84
Kalahari sands Northern 262 18.4 27 86

Griqualand West is classified as a semi-arid region (Fischer and Turner, 1978), with
an average annual rainfall ranging from 262 to 411 mm from west to east, most of which falls
in the summer months (Table 1). The low and erratic rainfall means that Griqualand West has
few perennial rivers. The most prominent is the Orange River, which originates in higher
rainfall areas to the southeast and forms the southern and south-western boundary of
Griqualand West. Many smaller rivers and tributaries in the region remain dry for many
months, or in extreme droughts, years at a time. The mean annual temperature of Griqualand
West is around 17–18 °C (Table 1). However, mean maximum and minimum temperatures of
higher lying areas and mountain ranges are cooler than surrounding lower-lying Kalahari
sands in both summer and in winter (Mucina and Rutherford, 2006). Thus, the mountain
ranges and, most prominently the elevated Ghaap Plateau, have a cooler climate.
The vegetation of Griqualand West can be broadly described as Savanna, specifically
forming part of the Eastern Kalahari Bushveld Bioregion (Mucina and Rutherford, 2006). The
Savanna Biome is characterized by an herbaceous ground layer dominated by forbs and
grasses, and a scattered upper layer comprising of woody vegetation. Some of the important
grass species in Griqualand West include Aristida canescens Henrard, A. congesta Roem. &
Schult., Brachiaria nigropedata (Ficalho & Hiern) Stapf, B. serrata (Thunb.) Stapf,
Cymbopogon pospischilli (K.Schum.) C.E.Hubb., Digitaria eriantha Steud., Enneapogon
cenchroides (Licht. ex Roem. & Schult.) C.E.Hubb., Eragrostis cylindriflora Hochst., E.
superba Peyr., Heteropogon contortus (L.) P.Beauv. ex Roem. & Schult., Melinis repens
(Willd.) Zizka and Themeda triandra Forssk. (Mucina and Rutherford, 2006). Important forb
species include Barleria macrostegia Nees, Dicoma capensis Less., Harpagophytum
procumbens (Burch.) DC. ex Meisn., Helichrysum cerastioides DC., Hermannia tomentosa
(Turcz.) Schinz ex Engl., Hermbstaedtia odorata (Burch. ex Moq.) T.Cooke, Hibiscus

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marlothianus K.Schum. and Jamesbrittenia aurantiaca (Burch.) Hilliard (Mucina and
Rutherford, 2006). Important woody species include Boscia albitrunca (Burch.) Gilg & &
Benedict, Dichrostachys cinerea (L.) Wight & Arn., Ehretia rigida (Thunb.) Druce, Euclea
crispa (Thunb.) Guerke, Grewia flava DC., Gymnosporia buxifolia (L.) Szyszyl., Olea europaea
L., Searsia lancea (L.f.) F.A.Barkley, Senegalia caffra (Thunb.) P.J.H. Hurter & Mabb., S.
mellifera (Benth.) Seigler & Ebinger, Tarchonanthus camphoratus L., Vachellia erioloba
(E.Mey.) P.J.H. Hurter, V. karroo (Hayne) Banfi & Galasso, V. tortilis (Forssk.) Banfi & Galasso
and Ziziphus mucronata Willd. (Mucina and Rutherford, 2006).

2.2 Assessment of endemicity

A list of plant taxa potentially endemic or near-endemic to the Griqualand West region
was compiled from various sources (Supplementary Table 1), including Wilman (1946),
Acocks (1988), Van Wyk and Smith (2001), Germishuizen and Meyer (2003), and Mucina and
Rutherford (2006). Additionally, all published volumes of the Flora of southern Africa (e.g.
Codd, 1985) were consulted to identify plant taxa with distributions centred in the Griqualand
West region. The resulting list comprised 85 taxa considered as potentially endemic to the
Griqualand West region. Distribution data of these species were supplemented with specimen
data housed in herbaria with substantial collections from the Griqualand West region, namely
A.P. Goossens Herbarium (PUC), Geo Potts Herbarium (BLFU), McGregor Museum
Herbarium (KMG), National Museum Herbarium (NMB), Pretoria National Herbarium (PRE)
and the South African National Parks Herbarium (KSAN). Fuzzy Gazetteer (Kohlschuetter,
2003) was used to clarify descriptive locality records. Information regarding the substrate and
geology on which specimens were collected was recorded from herbarium sheets.
Once all available locality data were collected for each of the potential endemic taxa,
the distribution of each species was mapped with ArcGIS (ESRI, 2011). GPS records were
limited and therefore the quarter-degree-grid (QDG: ca. 675 km2) system was often applied
for mapping purposes.
Assessing the resulting distribution maps and identifying taxa endemic to the
Griqualand West region was based on "Intuitive discernment" (Rosen, 1988). Species were
regarded as endemic if their distributions were centered (Fig. 1a) over the GWC boundaries
proposed by Van Wyk and Smith (2001). These boundaries were based on underlying geology
and used as a reference point for the position of the GWC. Distributions beyond these
boundaries in Griqualand-West did not disqualify endemicity. A near-endemic was defined as
a taxon with a distribution centered over the GWC boundaries proposed by Van Wyk and
Smith (2001), but with a population or populations disjunct in other centres of endemism (Fig.
1b). Floristic elements were regarded as taxa with a high density of locality records in the
Griqualand West region, but with sparse distributions elsewhere in southern Africa (Fig. 1c).

5
All taxa with distributions adjudged to not belong to one of the three afore-mentioned
categories were discarded (Fig. 1d).

a b)
)

c) d
)

Figure 1: Mapping criteria. a) Endemic: centered and restricted distribution in

Griqualand West; b) Near-endemic: distribution shared between Griqualand West
and another recognized centre of endemism (Gariep Centre in this case); c)
Floristic element: widespread with a clear clustering in Griqualand West, but
sparse occurrences beyond its boundaries, and d) Widespread and not endemic.

2.3 Mapping the GWC

The distributions of the endemic and near endemic taxa identified in section 2.2 were
overlaid using ArcGIS (ESRI, 2011) to create a quarter-degree-grid distribution map. All grids
containing at least one of the 21 endemic species were incorporated into the definition of the
GWC’s extent of occurrence. The outermost grids in all cardinal directions in which an endemic
occurs were taken as the border of the GWC. This border was smoothed to follow the geology
of the area and to reflect a more likely natural distribution of endemic species.

2.4 Comparison with other centres

Species lists were obtained for the quarter-degree grids (QDGs) in which the GWC is
located from BODATSA (Ranwashe, 2015). These lists were supplemented with herbarium
records obtained in section 2.2. The ratio between number of endemic taxa and total number
of species occurring in the GWC QDG’s was calculated to determine the level of endemism
for the centre. The relationship between total number of species comprising its flora, and the
number of endemic species was tested using R statistical software (R Core Team, 2016) for
all 18 southern African centres, by utilizing Spearman’s rank correlation coefficient. As there

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was a large correlation, it was possible to do a weighted regressions also using R statistical
software (R Core Team, 2016) for all 18 centres by comparing the total number of species
against the number of endemics. This was done for both the untransformed and the natural
log (ln) transformed data.

2.5 Core area concept

The GWC borders were further refined by excluding grids in which there were less than
three records of endemic species. Cross checks confirmed that where two or less endemics
were present in a grid that they occurred in areas bordering outlying geological features of the
GWC and that the majority of the grid would not harbor appropriate habitat. This exclusion of
grids allowed for more accurate demarcation of the core area of the GWC in which the density
of endemic taxa was the highest. Outer grids of the refined area were again taken as the outer
border of the GWC.

3. Results and Discussion

3.1 Floristic elements
The distributions of the identified floristic elements indicated links to both the Nama-
Karoo and Savanna Biomes. Eight taxa extend from GWC to the Nama Karoo Biome and are
as follows: Aizoon asbestinum Schltr. (succulent forb), Digitaria polyphylla Henrard (grass),
Eragrostis macrochlamys Pilg. var. wilmaniae (C.E.Hubb. & Schweick.) De Winter (grass),
Euphorbia wilmaniae Marloth (succulent forb), Indigofera damarana Merxm. & A.Schreib.
(forb), Pharnaceum viride Adamson (forb), Stachys burchelliana Launert (forb) and Titanopsis
calcarea (Marloth) Schwantes (succulent forb). The strongest link seems to lie in the Poaceae
and succulent forb species. This link may be explained by the relatively arid climate of the
Nama Karoo and the adaptability shown by these drought tolerant grasses and succulents to
such conditions in GWC (Blake and Jordan, 1993).
Sixteen floristic elements have shared distributions extending from GWC to the
broader Savanna Biome and are as follows: Carex burchelliana Boeck. (forb), Corchorus
pinnatipartitus Wild (forb), Crotalaria griquensis Bolus (forb), Ebracteola wilmaniae (L.Bolus)
Glen (forb), Erucastrum griquense (N.E.Br.) O.E.Schulz (forb), Euphorbia bergii A.C.White,
R.A.Dyer & B.Sloane (succulent forb), Euphorbia duseimata R.A.Dyer (succulent forb),
Helichrysum spiciforme DC. (forb), Ipomoea suffruticosa Burch. (forb), Jamesbrittenia albiflora
(Verd.) Hilliard (forb), Lotononis crumanina Benth. (forb), Pentzia oppositifolia Magee (forb),
Selago mixta Hilliard (forb), Sesbania notialis J.B.Gillett (forb), Sutera griquensis Hiern (forb)
and Triaspis hypericoides Burch. (forb). The floristic link is forb dominated by members of the
Fabaceae and Scrophulariaceae, and to a lesser extent by the Asteraceae and
Euphorbiaceae. The dominance of the Fabaceae is largely expected for areas containing

7
ironstone (Gibson et al., 2010) and it is also the most dominant family in the Savanna Biome
within which the GWC lies. The observed link with the Scophulariaceae may potentially be a
result of the complex geology of the region and the observed occurrence of members of this
family occurring as endemics in geologically defined centres (Allison and Stevens, 2001;
Siebert et al., 2001).

3.2 Endemic and near-endemic taxa

Of the 86 taxa that were potentially endemic to the GWC (Supplementary Table 1), 36
were discarded due to unfitting distributions, nomenclatural changes or a lack of data, and 26
taxa were found to be floristic elements of the GWC. Twenty-two taxa were found to be
endemic to the GWC, and two near-endemic. The distributions of the 22 endemics and two
near-endemics (Fig. 1, suppl. data) were roughly centered in the GWC boundaries of Van Wyk
and Smith (2001). These GWC endemics are as follows (arranged alphabetically by family,
then by species):

Acanthaceae
 Barleria media C.B.Clarke in Fl. Cap. 5(1): 51 (1901). Type: South Africa, Kalahari
region, Bechuanaland [Griqualand West], on the rocks at Chue Vley, Oct 1812,
Burchell 2386 (K, holo.). Perennial forb (0.05-0.1 m) assessed as Vulnerable
(Raimondo et al., 2009) and found on the Ghaap Plateau, Kuruman Hills and in river
valleys in the Northern Cape and North-West (Obermeyer, 1933).
 Blepharis marginata (Nees) C.B.Clarke in Fl. Cap. 5(1): 29 (1901); Acanthodium
marginatum Nees in Prodr. 11: 275 (1847). Type: South Africa, Hay division,
Griqualand West, Griqua Town [Griquatown], Burchell 1902 (G-DC, holo; K, PRE, iso.).
Dwarf shrub (0.05-0.1 m) found on the Asbestos Hills, Ghaap Plateau, Kuruman Hills,
Langberg and in river valleys in the Northern Cape (Vollesen, 2000).
 Glossochilus burchellii Nees., in Prodr. 11: 83 (1847). Type: Griqualand West, Hay
Division, plains between Griqua Town [Griquatown] and Witte Water, Burchell 1976.
Bechuanaland [Griqualand West]; near Kuruman, near the source of Kuruman
river, Burchell 2471 (K, syn.). Perennial forb (0.05-0.2 m) found on the Ghaap Plateau,
Kuruman Hills and Asbestos Hills in the Northern Cape (Clarke, 1901).
 Justicia puberula Immelman, FSA 30 (3:1): 33 (1995); Justicia parvibracteata
Immelman, Bothalia 16(1): 39 (1986). Type: South Africa, Cape Province [Northern
Cape], 11 miles NNW of Olifantshoek, in Toto Mountains, kloof, in rock crevices and
under shrubs, Tölken & Schlieben 1176 (PRE, holo.). Dwarf shrub (0.15-0.5 m) found

8
 on the Ghaap Plateau, Kuruman Hills, Langberg and in river valleys in the Northern
Cape (Immelman, 1995).
 Justicia thymifolia (Nees) C.B.Clarke in Fl. Cap. 5(1): 64 (1901); Adhatoda thymifolia
Nees in Prodr. 11: 392 (1847). Type: South Africa, Kalahari region, Hay division,
Griqualand West, between Griquatown and Spuigslang, Burchell 1702 (K, lecto.
designated by Immelman (1995); PRE photo). Shrub (0.35-1.75 m) found on the
Asbestos Hills, Ghaap Plateau, Kuruman Hills, Langberg and in river valleys in the
Northern Cape (Immelman, 1995).

Aizoaceae
 Antimima lawsonii (L.Bolus) H.E.K.Hartmann, Bothalia 28(1): 74 (1998);
Mesembryanthemum lawsonii L.Bolus, Ann. Bolus Herb. 4: 85 (1927). Ruschia
lawsonii (L.Bolus) L.Bolus, Notes Mesembryanthemum: 219 (1950). Type: South
Africa, Hay district, Papkuil, Aug 1912, Lawson 18551 (BOL, holo.). Perennial
succulent forb (0.1-0.25 m) found on the Asbestos Hills, Ghaap Plateau and Kuruman
Hills in the Northern Cape (Hartmann, 1998).
 Hereroa wilmaniae L.Bolus, Notes Mesembryanthemum 2: 82 (1929). Type: South
Africa, Griqualand West, near Dunmury, Oct 1922, Wilman 17264 (BOL, lect.
designated by Hartmann 2001). Prepodesma uncipetalum N.E.Br. in Gard. Chron.
3(89): 389 (1931); H. uncipetala (N.E.Br.) L.Bolus, Notes Mesembryanthemum 3: 135
(1938). Type: South Africa, Griekwaland West, Campbell, Pole Evans 6891 (K, holo.).
H. wilmaniae L.Bolus var. langebergenesis L.Bolus, Notes Mesembryanthemum 2: 82
(1929a). Type: South Africa, Langeberg [Langberg], Mar 1923, Wilman 17352 (BOL,
holo.). Perennial succulent forb (0.05-0.1 m) assessed as Rare (Raimondo et al., 2009)
and found on the Asbestos Hills, Ghaap Plateau, Kuruman Hills, Langberg and river
valleys in the Northern Cape (Hartmann, 2001).
 Lithops aucampiae L.Bolus subsp. euniceae (de Boer) D.T.Cole, Lithops: Flowering
Stones: 220 (1988). L. aucampiae L.Bolus var. euniceae de Boer in Succulenta 45(4):
491 (1966).Type: South Africa. Northern Cape, 13 km north of Hopetown, Cole 48
(PRE, holo.). L. aucampiae L.Bolus var. fluminalis D.T.Cole, Natl. Cact. Succ. J., 25:
8 (1970); L. aucampiae L.Bolus ssp. euniceae (de Boer) D.T.Cole var. fluminalis
D.T.Cole, Lithops: Flowering Stones: 220 (1988).Type: South Africa, Hopetown, 21
July 1968, Cole 54 (PRE, iso.). Perennial succulent forb (0.02 m) assessed as
Vulnerable (Raimondo et al., 2009) and found on the Asbestos Hills and river valleys
in the Northern Cape. (Hartmann, 2001).

9
  Lithops bromfieldii L.Bolus, Notes Mesembryanthemum 2: 452 (1934). Type: South
Africa, Gordonia division, Upington, Bromfield 2286/33 (BOL, holo.). L. glaudinae de
Boer, Succulenta: 129 (1960); L. bromfieldii L.Bolus var. glaudinae (de Boer) D.T.Cole,
Excelsa 3: 50 (1973). Type: South Africa, Glen Lyon, Matsap, Cole 116 (PRE). L.
insularis L.Bolus, Notes Mesembryanthemum 3: 75 (1937); L. bromfieldii L.Bolus var.
insularis (L.Bolus) B.Fearn, Cact. Succ. J. 42: 92 (1970). Type: South Africa, Kamies,
5 miles N of Orange River Bank, 23 miles SW of Upington Station, Wilmot 1353/34
(BOL, holo.). L. mennellii L.Bolus, Notes Mesembryanthemum 3: 75 (1937); L.
bromfieldii L.Bolus var. mennellii (L.Bolus) B.Fearn, Cact. Succ. J. 42:92 (1970). Type:
South Africa, Gordonia division [Griqualand West], Mennell 645/34 (BOL, holo.).
Perennial succulent forb (0.03 m) found in the Langberg region in the Northern Cape
(Hartmann, 2001).
 Lithops lesliei (N.E.Br.) N.E.Br. subsp. burchellii D.T.Cole, Lithops: Flowering
Stones: 217 (1988). Type: South Africa, Cape Province, Postmasburg, NE of Douglas,
in calcrete, Cole 302 (PRE, holo., iso.). Perennial succulent forb (0.01 m) assessed as
Near Threatened (Raimondo et al., 2009) and found on the Asbestos Hills and Ghaap
Plateau of the Northern Cape (Hartmann, 2001).
 Prepodesma orpenii (N.E.Br.) N.E.Br. in Gard. Chron. 3(89): 389 (1931).
Mesembryanthemum orpenii N.E.Br. Gard. Chron. 3(70): 303 (1921); Aloinopsis
orpenii (N.E.Br.) L.Bolus in S. African Gard. 19: 288 (1929); Nananthus orpenii
(N.E.Br.) L.Bolus, Notes Mesembryanthemum 3: 133 (1938). Type: South Africa,
Griqualand West, discovered by Mr Redmond Orpen at Campbell, Pole Evans 6910
(K, holo.). Rabiea tersa N.E.Br., Gard. Chron. 3(89): 54 (1931); Nananthus tarsus
(N.E.Br.) G.D.Rowley, Natl. Cact. Succ. J., 33(1): 6 (1978). Type: South Africa,
Northern Cape Province, Kuruman, Pole Evans 7623 (K, holo). Perennial succulent
forb (0.05 m) found on the Asbestos Hills, Ghaap Plateau, Kuruman Hills and Langberg
in the Northern Cape (Hartmann, 2001).

Anacardiaceae
 Searsia tridactyla (Burch.) Moffett in Bothalia, 37(2): 173 (2007); Rhus tridactyla
Burch., Trav. S. Africa, 1: 340 (1822). Type: South Africa, Northern Cape, Asbestos
Mountains, 27 Sep 1811, Burchell 1667 (K, holo.; BOL, iso.). Large shrub (0.5-4 m)
found on the Asbestos Hills, Ghaap Plateau, Kuruman Hills, Langberg and in river
valleys in the Northern Cape (Moffett, 1993).

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Asteraceae
 Amphiglossa tecta (Brusse) Koek. in Bothalia 29(1): 72 (1999); Pterothrix tecta
Brusse in Bothalia 20(1): 67 (1990). Type: South Africa, Northern Cape, Hay district,
Witsand [Witsand Nature Reserve], some 70 km SW of Postmasburg, farm Witsand
250, approximately 2 km W of Doornaar homestead, 26 Nov 1989, Brusse 5629 (PRE,
holo.; AD, B, BAF, BH, BM, BOL, BR, BRI, C, CAN, CANB, COI, E, EA, G, GH, GRA,
HBG, J, K, L, LD, LG, LISU, LMA, M, MEL, MO, NBG, NH, NSW, NU, O, P, R, S,
SRGH, U, UC, UPS, US, W, WAG, WIND, Z, iso.). Dwarf shrub (0.3-0.6 m) assessed
as Critically Rare (Raimondo et al., 2009), found along the Asbestos Hills, Langberg
and Ghaap Plateau in the Northern Cape (Koekemoer, 1999).
 Cineraria exilis DC. in Prodr. 6: 305 (1838) Type: South Africa, Vryburg division, at
the source of the Moshaweng River near Takun, 27 Sept 1812, Burchell 2274 (G-DC,
holo.; K, iso.). Perennial forb (0.15 m) found on the Ghaap Plateau and Kuruman Hills
in the Northern Cape and North-West (Cron et al., 2006).
 Dicoma kurumanii S.Ortiz & Netnou in Bot. J. Linn. Soc. 147(4), 510 (2005). Type:
South Africa, Northwest Province, Kuruman District, top of Ga Mhana Peak, 11 Feb
1886, Marloth 1103 (NBG, holo.; K, PRE, iso.). Dwarf shrub (0.5 m) found on the
Ghaap Plateau and Kuruman Hills in the Northern Cape (Ortiz and Netnou, 2005)
 Eriocephalus ericoides (L.f.) Druce subsp. griquensis M.A.N.Müll. in FSA 33(4:1):
49 (2001). Type: South Africa, Northern Cape, Herbert district, farm Eureka, Acocks
8753 (BOL, holo.; PRE, iso.). Kapokbos is a dwarf shrub (0.3-1 m) found on the
Asbestos Hills, Ghaap Plateau, Kuruman Hills, Langberg and in river valleys in the
Northern Cape (Müller et al., 2001).
 Gnaphalium englerianum (O.Hoffm.) Hilliard & B.L.Burtt in Bot. J. Linn. Soc. 82(3):
193 (1981); Amphidoxa engleriana O.Hoffm. in Bot. Jahrb. Syst. 10: 274 (1889). Type:
South Africa, Betschuanaland [Griqualand West], Kachun near Kuruman, 1200m, Feb
1886, Marloth 1004 (BOL, PRE, SAM, iso.). Perennial semi-decumbent forb (0.1 m)
found on the Ghaap Plateau, Kuruman Hills and in river valleys in the Northern Cape
(Hilliard and Burt, 1981; Hilliard, 1983).
 Pentzia stellata (P.P.J.Herman) Magee in Bot. J. Linn. Soc. 178(4): 644 (2015);
Rennera stellata P.P.J.Herman in Bot. J. Linn. Soc. 129(4): 368 (1999). Type: South
Africa, Northern Cape, Koopmansfontein Agricultural Research Station, 5 Mar 1998,
Herman 1482 (PRE, holo.; B, BM, C, E, J, G, K, KMG, M, MO, NBG, NMB, P, PRU, S,
UPS, WAG, WIND, Z, iso.). Perennial forb (0.5 m) assessed as Near Threatened
(Raimondo et al., 2009) and found on the Ghaap Plateau, primarily next to
unweathered calcrete pans in the Northern Cape and North-West (Herman, 1999).

11
  Tarchonanthus obovatus DC., Prodr. 5: 431 (1836). Type: South Africa,
Bechuanaland division [Griqualand West], Klipfontein, 19 Jun 1812, Burchell 2155 (G-
DC, holo.; GH, K, M, P, iso.). The Gordonia Camphor-Bush is a shrub (2 m) found on
the Asbestos Hills, Ghaap Plateau, Kuruman Hills, Langberg and in river valleys in the
Northern Cape (Herman, 2002).

Celastraceae
 Maytenus ilicina (Burch.) Loes., in Nat. Pflanzenfam., 2nd ed., 20b: 140 (1942);
Celastrus ilicinus, Burch. Trav. S. Africa 1: 340 (1822); Gymnosporia ilicina (Burch.)
Davidson in Bothalia 2: 296 (1927). Type: South Africa, Griekwaland West [Griqualand
West], Asbestos mountains, Burchell 1663 (K, holo.). The Gordonia Koko-Tree is a
large shrub (3 m) found on the Asbestos Hills, Ghaap Plateau, Kuruman Hills and
Langberg in the Northern Cape (Coates Palgrave, 2002).
 Putterlickia saxatilis (Burch.) Jordaan, in SAJB 64(6): 328 (1998); Celastrus saxatalis
Burch., Trav. S. Africa, 2: 264 (1824); Gymnosporia saxatalis Davidson in Bothalia 2:
303 (1927). Type: South Africa, Asbestos mountains, Griqua Town [Griquatown],
Burchell 1671 (K, holo.). The Gordonia False-spikethorn is a shrub (1.5-2 m) found on
the Asbestos Hills, Ghaap Plateau, Kuruman Hills, Langberg and valleys in the
Northern Cape (Jordaan and Van Wyk, 1998).

Fabaceae
 Calobota cuspidosa (Burch.) Boatwr. & B.-E.Van Wyk, in SAJB 75(3): 553 (2009);
Spartium cuspidosum Burch., Trav. S. Africa 1: 348 (1822). Type: South Africa,
Northern Cape, between ‘Gatikamma’ and ‘Klaarwater’ [Griquatown], Burchell 1697
(K, holo.). Lebeckia macrantha Harv. in Fl. Cap. 2: 83–84 (1862). Type: South Africa,
without precise locality, ‘Zooloo country’, Miss Owen s.n. (TCD, photo.). The
Griqualand Porcupine Bush is a shrub (1.5-4 m) found on the Asbestos Hills, Ghaap
Plateau, Kuruman Hills, Langberg and valleys in the Northern Cape (Boatwright et al.,
2018).

Poaceae
 Brachiaria dura Stapf. var. pilosa J.G.Anderson in Kirkia I: 104 (1960‒1961). Type:
South Africa, Hay division, Witsand [Witsand Nature Reserve], at foot of dunes, Apr
1940, Esterhuysen 2269 (PRE, holo.; K, iso.). Perennial grass (0.5-1.3 m) found in the
Langberg region in the Northern Cape (Fish et al. 2015).

12
Stilbaceae
 Nuxia gracilis Engl., in Bot. Jahrb. Syst. 10: 243 (1888); Lachnopylis gracilis (Engl.)
C.A.Smith, Bull. Misc. Inform. Kew 1930(1): 17 (1930). Type: South Africa, Hay
division, Klein Boetsap, Feb 1886, Marloth 980 (PRE, lecto. designated by
Leeuwenberg (1975); K, PRE iso.). The Sticky Nuxia is a shrub (1.5-4 m) found on the
Asbestos Hills, Ghaap Plateau, Kuruman Hills and valleys in the Northern Cape
(Leeuwenberg, 1975).

3.2 Phytogeographic patterns and geographical affinities of GWC endemics

Some of the GWC endemics are very rare, known from only a small region or a few
localities such as the forbs Amphiglossa tecta, Dicoma kurumanii and Gnaphalium
englerianum, and the cryptic succulents Lithops aucampiae subsp. euniceae and Lithops
lesiei subsp. burchellii (Fig. 1, suppl. data). There are also a number of GWC endemics that
are widely distributed throughout most of the centre. These specifically include the shrubs
Calobota cuspidosa, Putterlickia saxatilis, Searsia tridactyla and Tarchonanthus obovatus.
The two near-endemics, Justicia thymifolia and Nuxia gracilis, indicate links with two other
centrs of endemism, namely Gariep (Jürgens, 1991, Van Wyk and Smith, 2001) and
Sekhukhuneland (Siebert et al., 2002) respectively. In the GWC, the Asteraceae (seven
species) and Aizoaceae (six species) contributed the most endemics, with the Acanthaceae
(five species) also making a contribution with these three families contributing 18 of the 24
endemic and near-endemic taxa. The Asteraceae is diverse in southern Africa (McKenzie and
Barker, 2008) and one of the most dominant families in the GWC, possibly explaining the high
number of endemics associated with it. Dolomitic areas are often characterized by endemics
in the Asteraceae (Allison and Stevens, 2001). The large number of endemics in the
Aizoaceae may be partially explained by the arid nature of the centre and the succulence of
the endemics involved (Valente et al., 2013).
Only one of the 21 GWC endemics had a distribution that was restricted to the Nama-
Karoo Biome in the southern extremity of the centre (Prieska region), namely Lithops
aucampiae subsp. euniceae (Fig. 1, suppl. data). All the other endemics preferred the savanna
areas which covers more than 95% of the GWC. However, as this region in the Nama-Karoo
Biome was also host to numerous other GWC endemics, it was not excluded from the spatial
definition of the centre. Floristically it fits with the southern part of the Ghaap Plateau (Frisby,
2016).
In three other centres of endemism in the Savanna Biome of South Africa, the two
largest families with endemics were the Asphodelaceae and Asteraceae (Soutpansberg
Centre; Hahn, 2017), the Araceae and Vitaceae (Sekhukhuneland Centre; Siebert et al.,
2001), and the Asteraceae and Fabaceae (Barberton Centre; Williamson and Balkwill, 2015)

13
respectively. The Asteraceae is one of the largest endemic-bearing families in three of the four
Savanna centres of endemism, a trend that may be explained by the diversified nature of the
Asteraceae in both southern Africa and the world. However the other largest endemic bearing
families in four centres of endemism in the Savanna Biome are all different, indicating differing
selective pressures along different phylogenetic lineages (Kessler, 2002).
The GWC can broadly be subdivided into four floristic regions based on geology,
topography and the associated climatic variations (Frisby, 2016), namely the Ghaap Plateau
(GP), Ironstone Hills (IH; Kuruman and Asbestos Hills), the Langberg (LB), and the low lying
northern areas filled with Kalahari sands (KS). The substrate in the GWC which supports the
most endemic and near-endemic species is the soils derived from calcrete, dolomite and
limestone rocks of the GP. High biodiversity and endemism is globally well known for these
types of habitats (Clements et al., 2006). Thus, the flora of the GWC appears to be strongly
influenced by Ca- or Mg-rich substrates. This is evident when considering not only the
presence of GWC endemics, but also that several known calcicoles (species adapted to grow
on calcium-rich substrates) that are widespread in the arid regions of southern Africa, have
abundant distributions in the GWC, namely Bergia anagaloides, Eragrostis macrochlamys var.
wilmaniae, Erucrastrum griquense, Justicia puberula, Lotononis crumaniana and Salvia
namaensis (Van Wyk and Smith, 2001). Of the 24 endemic and near-endemic species, nine
show a specific preference to soils rich in calcium. Of these, Nuxia gracilis and Pentzia stellata
appear to be edaphic specialists, having only been recorded occurring on Ca-rich soils. The
former is a near-endemic known also from the Roossenekal-Steelpoort region in
Sekhukhuneland Centre of Endemism. This is a widely disjunct distribution (± 700 km)
between the two centers (Fig. 1, suppl. data), likely because of the presence of Ca-rich
substrates and arid conditions in both (Van Wyk and Smith, 2001). The taxonomic status of
the plants from Sekhukhuneland and the GWC, however, deserves closer scrutiny (see further
on). Pentzia stellata occurs solely next to weathered calcrete pans. Twelve other GWC
endemics also have part of their distributions occurring within the GP, suggesting that 20 of
the 24 endemic/near-endemic taxa prefer the cooler and more mesic conditions of the plateau.
Twenty-one of the 24 endemics were recorded from the IH adjacent to the GP. Such
patterns of endemism is well known for ironstone formations globally (Jacobi et al., 2007).
Kuruman Hills hosts 18 of the GWC endemics, as the climate is similarly to GP more favorable
than the rest of the GWC. Ironstone formations are also known for their specific plant-soil
associations (Gibson et al., 2010). Lithops aucampiae subsp. euniceae is currently only known
from the slightly drier Asbestos Hills in the southern part of the IH. Two GWC endemics have
distributions restricted to the LB, namely Lithops bromfieldii and Brachiaria dura var. pilosa.
Thirteen GWC endemics have at least part of their distributions within the LB. No GWC
endemics have distributions restricted to the KS. Thirteen GWC endemics do have at least

14
part of their distributions within the KS, although at low frequencies, probably as unfavorable
arid conditions prevail here compared to the other regions.
The large numbers of GWC endemics that are restricted to or show distributional
preference for the Ca-rich substrates of the GP indicate high levels of holo-endemism.
However, it is likely that some of the GWC endemics are also either paleoendemics or
neoendemics. The GWC near-endemic Nuxia gracilis is likely a paleoendemic. However,
closer inspection of type specimens of this species may result in taxonomic differentiation
between the two populations due to the large distance in between. The other GWC near-
endemic, Justicia thymifolia is likely to be a paleoendemic due to the close proximity of the
two populations.
The endemic shrubs tend to be distributed throughout the defined GWC, but
succulents tend to be range restricted (Table 2). However, Van Wyk and Smith (2001)
proposed that many plant taxa in the GWC may remain undescribed, largely due to the severe
lack of botanical collections that have been made in the region, emphasising the need to
further investigate the botanical diversity in Griqualand West to better understand the endemic
flora.

Table 2.
Breakdown of endemism per region according to total, restricted range and growth forms of GWC
endemics. The ironstone hills are subdivided into the northern Kuruman and southern Asbestos ranges.

Region Number of Restricted GWC Forbs Succulents Dwarf Shrubs

GWC endemics (5) (6) shrubs (5) (7)
endemics
Ghaap Plateau 21 1 forb 5 4 5 7
Ironstone Hills: 21 1 succulent 4 5 5 7
Kuruman hills 17 0 4 3 4 7
Asbestos hills 15 1 succulent 1 5 3 7
Kalahari sands 14 0 2 2 4 6
Langberg 14 1 succulent; 1 1 3 4 6
grass

3.3 Mapping the GWC boundary

Following the identification of the GWC endemic and near-endemic species, and
overlaying all their distributions, new borders for the GWC are proposed (Fig. 2) by fitting
boundaries to the outer grids in which GWC endemics were present. The resultant boundaries
of the GWC based on the endemic distributions differ from that of Van Wyk and Smith (2001).
The borders generated by the current study are significantly larger, having been extended

15
most noticeably south, south-west and east towards Kimberley (Fig. 2), as the proposed
endemics have a wider distribution than previously thought. The northern and eastern borders
predicted by Van Wyk and Smith (2001) largely coincide with those of this study. This
proposed region represents a noticeable transition from more elevated rocks of various
sedimentary origins, to lower-lying patches of deep aoelian sand based on the habitat
preferences of the GWC endemics. The expanded borders of the GWC proposed in this study
are not clearly linked to any ecological regions which differ from the regions included in the
GWC of Van Wyk and Smith (2001), barring the inclusion of some pockets of Nama-Karroo in
the southern regions.

3.4 Comparison with other centres

The large size and lower than predicted level of endemism (1.4% versus 2.2% of Van
Wyk and Smith (2001)) prompted a comparison between the total number of species and total
number of endemic species identified from the new GWC borders with that of other recognized
centres of plant endemism in southern Africa. As would be expected, a correlation (greater
than 0.5) was found for the variables total number of species and total number of endemic
species. The residual value for the GWC for the untransformed data (Fig. 3a) was not larger
than 2 and is therefore not an outlier value for number of endemic taxa expected per total
number of species on a 95% confidence interval. The residual value for the GWC for the
natural log transformed data (Fig. 3b) remained below 2 and therefore shows the GWC to not
be an outlier value for number of endemic taxa per total number of species on a 95%
confidence interval. This result suggests that despite the large size and limited numbers of
endemic species, the GWC suggested by this study falls within the current norms for

Figure 2: Combined distribution of GWC endemic and near-endemic species

(24 taxa). The green polygon demarcates the total distribution of the endemic
species. The red polygon represents the boundaries of the GWC as proposed
by Van Wyk & Smith (2001).

16
recognizing centres of plant endemism based on its total species pool. Indeed similar trends
were identified by Cowling and Hilton Taylor (1997) which showed a correlation between the
size of an areas species pool and endemism levels in southern Africa. This seems to be often
the case for centres of endemism in semi-arid savanna regions where endemism levels are
dependent on the size of the species pool and not the area.

Figure 3a: Weighted regression of endemism and flora for 18 centres of endemism (Van Wyk & Smith, 2001).
Albany Centre (AC); Barberton Centre (BC); Chimanimani-Nyanga Centre (CIC); Drakensberg Alpine Centre
(DAC); Gariep Centre (GC); Great Dyke Centre (GDC); Griqualand West Centre (GWC); Hantam-Roggeveld
Centre (HRC); Kamiesberg Centre (KBC); Kaokoveld Centre (KOC); Knersvlakte Centre (KVC); Little Karoo
Centre (LKC); Maputaland Centre (MC); Pondoland Centre (PC); Soutpansberg Centre (SC); Sekhukhuneland
Centre (SKC); Wolkberg Centre (WC); Worcester-Robertson Karoo Centre (WRKC).

Figure 3b: Weighted regression of endemism and flora for 18 centres of endemism (Van Wyk & Smith, 2001)
after natural log (ln) transformation. Albany Centre (AC); Barberton Centre (BC); Chimanimani-Nyanga Centre
(CIC); Drakensberg Alpine Centre (DAC); Gariep Centre (GC); Great Dyke Centre (GDC); Griqualand West
Centre (GWC); Hantam-Roggeveld Centre (HRC); Kamiesberg Centre (KBC); Kaokoveld Centre (KOC);
Knersvlakte Centre (KVC); Little Karoo Centre (LKC); Maputaland Centre (MC); Pondoland Centre (PC);
Soutpansberg Centre (SC); Sekhukhuneland Centre17 (SKC); Wolkberg Centre (WC); Worcester-Robertson
Karoo Centre (WRKC).
3.5 Core area of the GWC
The newly proposed borders of the GWC (Fig. 2) was found to cover a very extensive
area ─ larger than the borders proposed by Van Wyk and Smith (2001), and also harboring
comparatively less endemics. As there was a clear increase of endemics per grid from the
periphery to the core of the centre, it was decided to further explore the mapping of GWC to
highlight those areas richest in endemics. No endemics with restricted range distribution
occurred in the peripheral areas and were not excluded. The new GWC borders were therefore
refined further by excluding grids harboring low levels of endemism (Fig. 4). The concept of
‘core area’ proposed by this study may be of use when working with large centres with few
endemic species. The ‘core area’ of the GWC is based on high frequencies of records for the
22 endemic species. The core area coincides with the three most prominent geologies of the
region, including the banded ironstone hills, quartzitic outcrops and the dolomitic Ghaap
Plateau.

Figure 4: Core area of the GWC (blue-shaded polygon) based on the

occurrence of endemic species. Core area grids have more than three records
of endemic species. Green polygon represents the GWC boundaries based on
the occurrence of at least one GWC endemic plant species per quarter-degree-
grid.

4. Conclusion and future prospects

This study addressed the lack of knowledge regarding plant endemism in the
Griqualand West region, and provides support for the recognition of endemic plants from this
region. The endemic and near-endemic species accounted for 1.4% of the flora and their
restricted distributions demarcated the GWC beyond current definitions. The level of
endemism matches the trend found in other local centres of floristic endemism in southern

18
Africa. Thus, the GWC can justifiably be recognised as a centre of endemism. Many of the
GWC endemics show preference for Ca-rich substrates. As the GWC borders identified in this
study are extensive, covering a large geographical area, the more refined GWC core area was
demarcated to aid focussed conservation and future research efforts. This study proposes the
concept of ‘core area’ to identify an area that contains all the endemics and the highest
frequencies thereof. This will allow for focused conservation planning. Within the GWC core
area, the ranges of banded ironstone and the Ca-rich substrates of the Ghaap Plateau account
for >90% of the endemics. Future studies should identify the abiotic variables that influence
the flora of the GWC and thus the distribution of its endemics, especially plant-soil associations
(Gibson et al., 2010). This may provide insight into the finer-scale distributions of the endemic
plants, potentially allowing the identification of GWC sub-centres of endemism, further
directing conservation and research efforts which is much needed in these unique ecosystems
(Clements et al., 2006; Jacobi et al., 2007).

5. Acknowledgements
We thank the curators of various herbaria in central South Africa for giving us access
to their collections. Additional data was kindly provided by Braam van Wyk, Nanette van
Staden, and Norbert Hahn. The financial assistance of the National Research Foundation
(Grant UID: 103370) towards this research is hereby acknowledged. Opinions expressed and
conclusions arrived at are those of the author and are not necessarily to be attributed to the
NRF.

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D.A., Manyama, P.A., 2009. Red List of South African Plants. Strelitzia 25. South African
National Biodiversity, Pretoria.
Ranwashe, F. 2015. BODATSA: Botanical Collections. v1.1. South African National
Biodiversity Institute. Dataset/Occurrence.
http://ipt.sanbi.org.za/iptsanbi/resource?r=brahms_online&v=1.1.
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Valente, L.M., Britton, A.W., Powell, M.P., Papadopulos, A.S., Burgoyne, P.M., Savolainen,
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Appendix. Supplementary data.
Distribution data

Supplementary Table 1
Taxa considered during the screening process for GWC endemics and near-endemics.
Justifications are based on herbarium records as were available on 30 January 2017.

Species Family Classification Justification

Distribution extends as far
Floristic
Agrostis griquensis Stapf Poaceae east as Bloemfontein;
element
centred in GWC.
Wide distribution extending
Aizoon asbestinum Schltr. Aizoaceae Discarded from GWC to southern
Namibia.
Amphiglossa tecta (Brusse) Koekemoer One locality at Witsand in
Asteraceae GWC endemic
Syn.: Pterothrix tecta Brusse western GWC.
Antimima lawsonii (L.Bolus)
Restricted to GWC, south
H.E.K.Hartmann Aizoaceae GWC endemic
and north of Lime Acres.
Syn.: Ruschia lawsonii (L.Bolus)
Restricted to GWC between
Barleria media C.B.Clarke Acanthaceae GWC endemic
Kuruman and Pomfret.
Bergia anagaloides E.Mey. ex Fenzl Widespread in South Africa
Elatinaceae Discarded
Syn.: Bergia alsinoides Friedr.-Holzh. and Namibia.
Restricted to central GWC
Blepharis marginata (Nees) C.B.Clarke
Acanthaceae GWC endemic between Sishen, Kimberley
Syn.: Acanthodium marginatum Nees
and south of Postmasburg.
Boscia foetida Schinz subsp. foetida Widespread in Gariep
Capparaceae Discarded
Syn.: Boscia rautanenii Schinz region and Namibia.
Brachiaria dura Stapf var. pilosa Restricted to GWC south of
Poaceae GWC endemic
J.G.Anderson Postmasburg.
Occurs too far east of
Brachystelma canum R.A.Dyer Apocynaceae Discarded
GWC; south of Mafikeng.
Occurs too far east of GWC
Brachystelma dimorphum R.A.Dyer
Apocynaceae Discarded in Free State near
subsp. dimorphum
Bloemfontein.
Calobota cuspidosa (Burch.) Boatwr. &
Restricted to GWC, but
B.-E.Van Wyk Fabaceae GWC endemic
widespread within it.
Syn.: Lebeckia macrantha Harv.
Calobota psiloloba (E.Mey.) Boatwr. & B.-
Not present in GWC,
E.Van Wyk Fabaceae Discarded
occurring in Eastern Cape.
Syn.: Lebeckia psiloloba (E.Mey.) Walp.

24
 Occurs as far east as
Floristic
Carex burchelliana Boeckeler Cyperaceae Klerksdorp. Centred in
element
GWC.
Restricted to GWC; known
Cineraria exilis DC. Asteraceae GWC endemic only from type specimen
found east of Kuruman.
Widespread in central
Convolvulus boedeckerianus Peter Convolvulaceae Discarded
South Africa.
Floristic Also occurs in southern
Corchorus pinnatipartitus Wild Malvaceae
element Botswana; centred in GWC.
Floristic Wide distribution in central
Crotalaria griquensis L.Bolus Fabaceae
element South Africa.
Restricted to GWC around
Dicoma kurumanii S.Ortiz & Netnou Asteraceae GWC endemic
Kuruman.
Distribution extends west as
Floristic
Digitaria polyphylla Henrard Poaceae far as Hoopstad; centred in
element
GWC.
Ebracteola wilmaniae (L.Bolus) Glen Distribution extends east to
Floristic
Syn.: Ruschia wilmaniae (L.Bolus) Aizoaceae just north of Bloemhof;
element
L.Bolus var. wilmaniae centred in GWC.
Eragrostis macrochlamys Pilg. var.
Also present in central
wilmaniae (C.E.Hubb. & Schweick.) De Floristic
Poaceae Namibia and western
Winter element
Botswana; centred in GWC.
Syn.: Eragrostis macrochlamys Pilg. p.p.
Eriocephalus ericoides (L.f.) Druce subsp.
griquensis M.A.N.Müll. Restricted to GWC, but
Asteraceae GWC endemic
Syn.: Eriocephalus ericoides (L.f.) Druce widespread within it.
p.p.
Widespread in central and
Erucastrum griquense (N.E.Br.)
north-western South Africa,
O.E.Schulz Floristic
Brassicaceae extending as far as
Syn.: Diplotaxis griquensis (N.E.Br.) element
Zimbabwean border;
Sprague
centred in GWC.
Extends too far into Free
Euphorbia bergii A.C.White, R.A.Dyer & Floristic State to south of
Euphorbiaceae
B.Sloane element Koffiefontein; centred in
GWC.
Floristic Also in Namibia; centred in
Euphorbia duseimata R.A.Dyer Euphorbiaceae
element GWC.
Euphorbia inornata N.E.Br. Sunken into E. crassipes
Euphorbiaceae Discarded
Syn. of Euphorbia crassipes Marloth Marloth.
Euphorbia planiceps A.C.White, R.A.Dyer
Sunken into E. wilmaniae
& B.Sloane Euphorbiaceae Discarded
Marloth.
Syn.: Euphorbia wilmaniae Marloth

25
Euphorbia rectirama N.E.Br. Sunken into E. spartaria
Euphorbiaceae Discarded
Syn.: Euphorbia spartaria N.E.Br. N.E.Br.
Euphorbia wilmaniae Marloth
Floristic Wide distribution in South
Syn.: Euphorbia planiceps A.C.White, Euphorbiaceae
element Africa; centred in GWC.
R.A.Dyer & B.Sloane
Wide distribution in South
Galenia portulacacea Fenzl Aizoaceae Discarded
Africa.
Restricted to GWC, but
Glossochilus burchellii Nees Acanthaceae GWC endemic
widespread within it.
Gnaphalium englerianum (O.Hoffm.)
Restricted to GWC around
Hilliard & B.L.Burtt Asteraceae GWC endemic
Reivilo.
Syn.: Amphidoxa engleriana O.Hoffm.
Halopeplis amplexicaulis (Vahl) Ung.-
Chenopodiaceae Discarded Cosmopolitan species.
Sternb. ex Ces., Pass. & Gibelli
Floristic Also present in southern
Helichrysum spiciforme DC. Asteraceae
element Namibia; centred in GWC.
Hereroa wilmaniae L.Bolus
Restricted to GWC, but
Syn.: Hereroa uncipetala (N.E.Br.) Aizoaceae GWC endemic
widespread within it.
L.Bolus
Indigofera damarana Merxm. &
A.Schreib. Floristic Also occurs in Namibia;
Fabaceae
Syn.: Indigofera wilmaniae Baker f. ex element centred in GWC.
J.B.Gillett
Ipomoea suffruticosa Burch. Widely distributed in
Floristic
Syn.: Turbina suffruticosa (Burch.) Convolvulaceae southern Africa; centred in
element
A.Meeuse GWC.
Distribution extends too far
Jamesbrittenia albiflora (I.Verd.) Hilliard Floristic east and south into Free
Scrophulariaceae
Syn.: Sutera albiflora I.Verd. element State; partially centred in
GWC.
Justicia puberula (Immelman) Immelman Restricted to GWC, but
Acanthaceae GWC endemic
Syn.: Justicia parvibracteata Immelman widespread within it.
Main population restricted
Justicia thymifolia (Nees) C.B.Clarke GWC near- to GWC and widespread
Acanthaceae
Syn.: Adhatoda thymifolia Nees endemic within it; marginally present
in Gariep Centre.
Listia minima (B.-E.van Wyk) B.-E.van
Fabaceae Discarded Unresolved taxon.
Wyk & Boatwr.
Distribution extends too far
Listia subulata (B.-E.van Wyk) B.-E.van Floristic
Fabaceae east into Free State;
Wyk & Boatwr. element
partially centred in GWC.
Lithops aucampiae subsp. aucampiae Variety no longer
Aizoaceae Discarded
var. aucampiae (de Boer) D.T.Cole recognised.

26
Lithops aucampiae subsp. aucampiae Variety no longer
Aizoaceae Discarded
var. koelemanii (de Boer) D.T.Cole recognised.
Lithops aucampiae L.Bolus subsp. Restricted to GWC between
Aizoaceae GWC endemic
euniceae (de Boer) D.T.Cole Hopetown and Douglas.
Lithops bromfieldi L.Bolus
Restricted to GWC south-
Syn.: Lithops bromfieldii L.Bolus var. Aizoaceae GWC endemic
west of Postmasburg.
glaudinae (de Boer) D.T.Cole
Lithops lesliei (N.E.Br.) N.E.Br. subsp.
Restricted to GWC between
burchellii D.T.Cole Aizoaceae GWC endemic
Griquatown and Campbell.
Syn.: Lithops lesliei (N.E.Br.) N.E.Br. p.p.
Floristic Widespread; centred in
Lotononis burchellii Benth. Fabaceae
element GWC.
Wide distribution in central
Floristic
Lotononis crumanina Burch. ex Benth. Fabaceae South Africa; centred in
element
GWC.
Distribution in Namibia
unknown; should be
Lotononis linearifolia B.-E.van Wyk Fabaceae Discarded investigated further as a
potential near-endemic
taxon.
Maytenus ilicina (Burch.) Loes.
Restricted to GWC, but
Syn.: Gymnosporia ilicina (Burch.) Celastraceae GWC endemic
widespread within it.
Davison
Marginally present in GWC,
Melhania transvaalensis Szyszyl. Malvaceae Discarded centred in Limpopo
Province.
Present in Eastern Cape,
Floristic Northern Cape and Free
Mestoklema copiosum N.E.Br. ex Glen Aizoaceae
element State provinces; centred in
GWC.
Monechma districhotrichum (Lindau)
Present in southern
P.G.Mey. Acanthaceae Discarded
Namibia.
Syn.: Justicia distichotrichum Lindau
Nananthus aloides (Haw.) Schwantes
Widespread in central
Syn.: Aloinopsis aloides (Haw.) Aizoaceae Discarded
South Africa.
Schwantes
Centred in North-West
Nerine frithii L.Bolus Amaryllidaceae Discarded Province; marginally
present in GWC.
Present too far west of
Nerine hesseoides L.Bolus Amaryllidaceae Discarded
GWC near Kroonstad.
Present in central and
Nolletia annetjieae P.P.J.Herman Asteraceae Discarded eastern Namibia and
southern Botswana.

27
 Present in GWC between
Nuxia gracilis Engl. GWC near- Vryburg and Douglas with a
Buddlejaceae
Syn.: Lachnopylis gracilis (Engl.) C.A.Sm. endemic disjunct population in
Sekhukhuneland Centre.
Restricted to GWC between
Pentzia stellata (P.P.J.Herman) Magee Asteraceae GWC endemic Vryburg and south of
Danielskuil.
Present too far east of
Floristic
Pentzia oppositifolia Magee Asteraceae GWC as far as Boshof;
element
centred in GWC.
Petalidium parvifolium C.B.Clarke ex
Mainly in Gariep region
Schinz Acanthaceae Discarded
including southern Namibia.
Syn.: Petalidium wilmaniae Oberm.
Also present in
Floristic Soutpansperg (Limpopo),
Pharnaceum viride Adamson Molluginaceae
element Western Cape and Gariep
region; centred in GWC.
Sunken into
Mesembryanthemum
amabile (Gerbaulet &
Phyllobolus amabilis Gerbaulet & Struck Aizoaceae Discarded
Struck) Klak
which also occurs in the
Western Cape.
Prepodesma orpenii (N.E.Br.) N.E.Br. Restricted to GWC, but
Aizoaceae GWC endemic
Syn.: Nananthus orpenii N.E.Br. widespread within it.
Putterlickia pyracantha (L.) Szyszyl.
Widespread in southern
Syn.: Catha campestris (Eckl. & Zeyh.) Celastraceae Discarded
Africa.
C.Presl
Putterlickia saxatilis (Burch.) M.Jordaan
Restricted to GWC, but
Syn.: Gymnosporia saxatilis (Burch.) Celastraceae GWC endemic
widespread within it.
Davison
Ruschia griquensis (L.Bolus) Schwantes
Syn.: Mesembryanthemum griquense Aizoaceae Discarded Widespread in South Africa.
L.Bolus
Sunken into more widely
Salsola atrata Botsch.
Amaranthaceae Discarded distributed Caroxylon
Syn.: Salsola globulifera Fenzl
atratum Botsch.
Distribution extends south
Floristic
Salsola humifusa A.Brückn. Chenopodiaceae and east, as far south as De
element
Aar; centred in GWC.
Widespread in central
Salvia namaensis Schinz South Africa and Western
Lamiaceae Discarded
Syn.: Salvia burchellii N.E.Br. Cape, and southern
Namibia.

28
Searsia tridactyla (Burch.) Moffett Restricted to GWC, but
Anacardiaceae GWC endemic
Syn.: Rhus tridactyla Burch. widespread within it.
Distribution extends too far
Selago mixta Hilliard Floristic into North-West Province,
Scrophulariaceae
Syn.: Walafrida paniculata (Thunb.) Rolfe element as far east as Rustenburg;
centred in GWC.
Septulina ovalis (E.Mey. ex Harv.) Tiegh. Not present in GWC; only in
Loranthaceae Discarded
Syn.: Loranthus ovalis E.Mey. ex Harv. Gariep Centre.
Distribution extends too far
Floristic
Sesbania notialis J.B.Gillett Fabaceae to the east, as far as
element
Bothaville; centred in GWC.
Present in southern
Stachys burchelliana Launert Floristic
Lamiaceae Namibia (outside of Gariep
Syn.: Stachys burchellii Benth. element
Centre); centred in GWC.
Stapelia hursita L. var. gariepensis
Only present in Gariep
(Pillans) Bruyns Apocynaceae Discarded
region.
Syn.: Stapelia gariepensis Pillans
Not present in GWC, only in
Stapelia rubiginosa Nel Apocynaceae Discarded
Gariep Centre.
Stapelia similis N.E.Br.
Apocynaceae Discarded Not present in GWC.
Syn.: Stapelia juttae Dinter
Also present in central
Sutera griquensis Hiern Floristic Free-State and central
Scrophulariaceae
Syn.: Sutera burchellii Hiern element North-West Provinces;
centred in GWC.
Restricted to GWC, but
Tarchonanthus obovatus DC. Asteraceae GWC endemic
widespread within it.
Sunken into widely
Thesium dumale N.E.Br. Santalaceae Discarded distribued T. resedoides
A.W.Hill
Titanopsis calcarea (Marloth) Schwantes Floristic Widely distributed in central
Aizoaceae
Syn.: Titanopsis fulleri Tischer element GWC.
Trachyandra burkei (Baker) Oberm. Also present in southern
Asphodelaceae Discarded
Syn.: Anthericum burkei Baker Botswana.
Also present in Limpopo,
Triaspis hypericoides (DC.) Burch. subsp. Floristic
Malpighiaceae Gauteng and Mpumalanga;
hypericoides element
centred in GWC.

29
a) b)

c) d)

e) f)

g) h)

i) j)

30
k) l)

m) n)

o) p)

q) r)

s) t)

31
 u) v)

w) x)

Figure 1: Global distribution of a) Amphiglossa tecta; b) Antimima lawsonii; c)

Barleria media; d) Blepharis marginata; e) Brachiaria dura var. pilosa; f) Calobota
cuspidosa; g) Cineraria exilis; h) Dicoma kurumanii; i) Eriocephalus ericoides
subsp. griquensis; j) Glossochilus burchelli; k) Gnaphalium englerianum; l)
Hereroa wilmaniae; m) Justicia puberula (near endemic); n) Justicia thymifolia; o)
Lithops aucampiae subsp. euniceae; p) Lithops bromfieldii; q) Lithops lesliei
subsp. burchellii; r) Maytenus ilicina; s) Nuxia gracilis (near endemic); t) Pentzia
stellata; u) Prepodesma orpenii; v) Putterlickia saxatilis; w) Searsia tridactyla; and
x) Tarchonanthus obovatus.

32