ArticlePDF Available

A new species of Polygala (Polygalaceae) from ultramafic soils in Sekhukhuneland, South Africa, with notes on its ecology

Authors:
  • North-West University, Mafikeng campus

Abstract and Figures

Polygala sekhukhuniensis Retief, Siebert & A.E.Van Wyk (Polygala; section Polygala; subsection Heterolophus), a new species with a restricted range in Sekhukhuneland, South Africa, is described, illustrated and compared with other members of the genus. It is a dwarf shrub that can be distinguished by its much-branched habit, sparsely flowered inflorescences, pink alae with darker pink veins, brown to black seed testa, and oblate pollen grains with pronounced opercula. Geographically, P. sekhukhuniensis is confined to heavily eroded localized sites, a natural geomorphological feature of some of the highly water-dispersible soils derived from ultramafic rocks in the valleys of the Steelpoort River and its tributaries in the Sekhukhuneland Centre of Plant Endemism. P. sekhukhuniensis is a calciotrophic excluder of heavy metals that accumulates Ca in its leaves. It is ecologically compared with co-occurring species of Polygala on ultramafic-derived soil.
Content may be subject to copyright.
A new species of Polygala (Polygalaceae) from ultramafic soils in
Sekhukhuneland, South Africa, with notes on its ecology
S.J. Siebert
a,
, E. Retief
b
, A.E. Van Wyk
c
, M. Struwig
a
a
A.P. Goossens Herbarium, School of Environmental Sciences and Development, North-West University, Private Bag X6001, Potchefstroom 2520, South Africa
b
National Herbarium, South African National Biodiversity Institute, Private Bag X101, Pretoria 0001, South Africa
c
H.G.W.J. Schweickerdt Herbarium, Department of Plant Science, University of Pretoria, Pretoria 0002, South Africa
Received 14 August 2009; received in revised form 7 January 2010; accepted 14 January 2010
Abstract
Polygala sekhukhuniensis Retief, Siebert & A.E.Van Wyk (Polygala; section Polygala; subsection Heterolophus), a new species with a
restricted range in Sekhukhuneland, South Africa, is described, illustrated and compared with other members of the genus. It is a dwarf shrub that
can be distinguished by its much-branched habit, sparsely flowered inflorescences, pink alae with darker pink veins, brown to black seed testa, and
oblate pollen grains with pronounced opercula. Geographically, P. sekhukhuniensis is confined to heavily eroded localized sites, a natural
geomorphological feature of some of the highly water-dispersible soils derived from ultramafic rocks in the valleys of the Steelpoort River and its
tributaries in the Sekhukhuneland Centre of Plant Endemism. P. sekhukhuniensis is a calciotrophic excluder of heavy metals that accumulates Ca
in its leaves. It is ecologically compared with co-occurring species of Polygala on ultramafic-derived soil.
© 2010 SAAB. Published by Elsevier B.V. All rights reserved.
Keywords: Bushveld Complex; Caruncle; Endemism; Heavy metals; Palynology; Sekhukhuneland Centre of Endemism; Taxonomy; Ultramafic rocks
1. Introduction
Since the pioneering work of Wild (1965) on plants
associated with soils rich in heavy metals and derived from
ultramafic rocks in southern Africa, and subsequent ground-
breaking research on plant-soil associations on southern African
serpentinites by Morrey et al. (1989), Williamson et al. (1997)
and Balkwill and Campbell-Young (2001), many plant species
new to science have been described from such substrates. In
recent years the Sekhukhune land Centre of Plant Endemism
(SCPE) in South Africa has been highlighted as an area
harbouring potential new plant species (Van Wyk and Smith,
2001). Siebert et al. (2001) showed that the high levels of plant
endemism of this region are significantly correlated with the
heavy metal soils derived from ultramafic rocks. In the last three
years, six new endemic plant species (including a new
monotypic genus, Prototulbaghia) were described from the
SCPE (Venter et al., 2007; Vosa, 2007; Burrows and Burrows,
2008; Hankey et al., 2008; Retief et al., 2008; Manning and
Goldblatt, 2009).
Massoura et al. (2004) have shown that plant species adapted
to heavy metal soils often accumulate or exclude metals that are
present in the soil at high concentrations. This physiological
adaptation is usually also expressed in the morphology through
ecological speciation (Balkwill and Campbell-Young, 2001;
Retief et al., 2008 ). Subsequent studies on heavy metal
accumulation by plants in the SCPE by Mandiwana et al.
(2007) have revealed several metal accumulators and excluders.
Such species are known to be genetically different from their
widespread congeners (Yang et al., 2005). This has warranted
taxonomic investigation into one such excluder that is tolerant of
soils rich in heavy metals, a member of Polygala that was initially
considered an ecotype of P. leptophylla Burch. var. leptophylla.
However, subsequent field work and comparative morphological
studies have shown the putative ecotype to be a distinct new
species related to members of section Polygala,subsection He-
terolophus (Paiva, 1998), specifically P. hottentotta Presl.,
P. leptophylla var. leptophylla and P. seminuda Harv. It is
confined to areas of highly water-dispersible soils derived from
A
vailable online at www.sciencedirect.com
South African Journal of Botany 76 (2010) 345 353
www.elsevier.com/locate/sajb
Corresponding author.
E-mail address: stefan.siebert@nwu.ac.za (S.J. Siebert).
0254-6299/$ - see front matter © 2010 SAAB. Published by Elsevier B.V. All rights reserved.
doi:10.1016/j.sajb.2010.01.003
ultramafic rocks. The new species is here described as Polygala
sekhukhuniensis Retief, Siebert & A.E.Van Wyk. This is the
second excluder of heavy metals, after Euclea sekhukhuniensis
(Retief et al., 2008), that is strictly confined to localized areas of
naturally eroded, ultramafic soils in Sekhukhuneland.
The aim of this paper is to describe and name the new taxon, and
highlight the morphological, palynological and ecological differ-
ences between the taxon and its most closely related congeners.
2. Materials and methods
2.1. Morphological assessment
Live material of the new species was extensively studied in the
field. Its morphology was compared to existing descriptions and
treatments of the genus (Exell, 1960; Paiva, 1998). Specimens of
Polygala housed in the National Herbarium (PRE), Pretoria and
H.G.W.J. Schweickerdt Herbarium (PRU), University of Pretoria,
were examined to gather quantitative and qualitative data on
morphology, phenology and distribution of the new species, as
well as related taxa. Micrographs of seed were taken with a Nikon
Digital Camera DXM 1200 F fitted on a Nikon SMZ 1500
stereomicroscope. Seed terminology follows Paiva (1998).
2.2. Palynological assessment
Mature, unopened flower buds from herbarium specimens
were dissected to obtain anthers with pollen. Unacetolyzed
pollen grains were mounted onto aluminium stubs, sputter-
coated with gold/palladium (Au/Pd) and examined with a FEI
Quanta 200 ESEM Scanning Electron Microscope (SEM). Ten
pollen grains per species were measured for polar length (P) and
equatorial width (E). Palynological terminology follows Punt et
al. (1994).
2.3. Plant-soil assessment
Plant material of the new species and P. hottentotta, as well
as soil samples were collected during early summer, environ-
mental factors were noted and associated plant communities
identified (Siebert et al., 2002). Samples were taken at ten sites,
five each dominated by either species. Soil analyses was done
with X-Ray Fluorescence (XRF) Spectrometry and plant
analyses with Atomic Absorption Spectrophotometry (AAS)
as well as Inductively Coupled Plasma-Mass Spectrometry
(ICP-MS). Terminology desc ribi ng exclus ion mechan isms
follows Massoura et al. (2004).
3. Taxonomy
The subcosmopolitan family, Polygalaceae, comprises a bout
19 genera and 925 species (Mabberley, 1997). The flowers are
superficially similar to those of members of the subfamily
Papilionoideae (Fabaceae). However, the wings (alae) of Po-
lygala flowers belong to the calyx and not to the corolla. The
standard is completely different and Polygala has a crest at the
tip of the keel. Currently various studies (Prenn er, 2004; Banks
et al., 2008) confirm the placin g of the erstw hile order
Polygalales in an expanded order Fabales (APG II, 2003). In
the Flora of southern Africa region, the genus Polygala is
represented by ± 88 species (Bredenkamp, 2000), of which ten
are currently known from the SCPE.
Polygala sekhukhuniensis Retief, Siebert & A.E.Van Wyk, sp.
nov.; fruticulus pumilus caulibus patentibus caudice lignoso;
caules foliaque sparse apprese hirsuti; folia spiraliter disposita,
lamine linearo-oblonge vel obovata, apice mucronata; calyx
sepalis duobus aliformibus, alis roseis; corolla carina cristata,
crista atrorosea vel purpurea.
Type. 2430 (Pilgrim's Rest): Mpumalanga, Thornecliffe
Chrome Mine, turn off from Lydenburg-Sekhukhune road, [1
December 1997], Van Wyk 13031 (PRU, holo.; PRE, iso.).
Rigid perennial shrublet, 0.20.4 m high, with a woody
rootstock; stems and leaves sparsely adpressed-hairy; hairs
unbranched. Stems usually more than four from rootstock, erect
and spreading, branched, woody at base; young branches green,
herbaceous. Leaves spirally arranged, shortly petiolate; petiole
up to 0.75 mm long; blade linear-oblong or obovate, 715× 1.0
2.5 mm, apex obtuse, occasionally acute, mucronate, margin
entire. Inflorescences comprising terminal racemes, peduncle up
to 80 mm long, pedicels 1.75 mm long. Flowers
irregular;
bi
sexual, drooping. Calyx pentamerous, with sepals unequal,
three outer ones sepaloid, two inner ones petaloid, larger wing-
like, 68 × 3.55.0 mm, all free; wings pink with darker pink
veins. Corolla of five petals, lowest one forming a carina, crested,
crest dark pink to purple, two lateral ones usually vestigial or
absent, two upper ones joined at base to carina and staminal tube.
Stamens eight, basally fused for ±two-thirds their length. Ovary
superior, two locular. Fruit a laterally compressed capsule, up to
6.5 mm long, membranous, edges wing-like, 0.25 mm wide,
deeply notched at apex, 2-seeded. Seed 4.5×1.5 mm, ellipsoid,
caruncle oblique, appendages poorly developed, single one
membranous, paired ones chitinous; indumentum silky white,
±1 mm longer than seed; testa dark-brown to black. Flowering
time: November to June. (Fig. 1).
3.1. Diagnostic characters and key to species
Polygala sekhukhuniensis and two related species, P.
hottentotta and P. leptophylla, are distinguished from P.
seminuda and the rest of the genus by their narrowly ovate
leaves, free anterior sepals, terminal racemes, wings not more than
6 mm broad, and the carina 6 mm long with crest 13mmlong.
P. sekhukhuniensis and P. leptophylla differ from P. hottentotta,
with the latter one having longer peduncles (124 mm versus 78
90 mm), greater number of flowers per raceme (23 versus 712),
larger flowers (wing length 7.8 mm versus 6.87.0 mm), and
smaller seed (4.3×1.5 mm versus 5.65.9× 1.9 mm) (Table 1).
Morphologically P. sekhukhuniensis most closely resembles
P. leptophylla var. leptophylla , however, the leaf size of the
former is 715 × 12.5 mm, whereas the leaves of the latter one
are 1422 × 1.53mm(Table 1). Flowers of P. sekhukhuniensis
differ from P. leptophylla var. leptophylla in the crest of the
former being purple and the alae pink with darker pink veins,
whereas the crest is white, pink or purple in the latter, with the
346 S.J. Siebert et al. / South African Journal of Botany 76 (2010) 345353
alae whi tish to yellowish with green veins. Seeds of P.
sekhukhuniensis differ from P. leptophylla var. leptophylla in
the colour of the testa, dark brown-black versus reddish-brown,
the bare patch on the testa between the caruncle and the
indumentum (the size, 0.3 × 0.6 mm versus 1.2 × 1.5 mm), and
the caruncular appendages, poorly developed (0.2 versus
0.8 mm long) (Fig. 2 ; Table 1).
Key to the species of Polygala in Sekhukhuneland
1a. Plants procumbent, up to 50 mm high P. krumanina
1b. Plants erect, 50 mm to 1.5 m high 2
2a. Anterior sepals united for at least half their length P. uncinata
2b. Anterior sepals free or only slightly joined at the base 3
3a. Flowers solitary or in lateral racemes 4
3b. Flowers in terminal racemes, sometimes with additional
lateral racemes 5
4a. Flowers presented in a raceme, 30140 mm long P. sphenoptera
4b. Flowers solitary, borne in leaf axils P. gerrardii
5a. Wings 715 mm broad; sturdy, erect shrublets up to 2 m tall P. virgata
5b. Wings less than 6 mm broad; slender, annual or perennial
herbs up to 0.5 m tall 6
(continued on next page)
Table 1
Mean measurements of selected morphological c haracters in Polygala
sekhukhuniensis and three closely related species (n =10/species).
Polygala
sekhukhuniensis
Polygala
leptophylla
Polygala
hottentotta
Polygala
seminuda
Number of flowers
per raceme
7±1.3 12 ±1.3 23 ±7.4 11 ±2.3
Inflorescence (mm)
Peduncle length 78 ±23 90 ±18 124±27 86 ±24
Flowers (mm)
Wing length 6.8±0.6 7.0±0.8 7.8±0.8 6.7±1.1
Wing breadth 4.1±0.7 3.5±0.5 4.0±0.7 3.7±0.9
Leaves (mm)
Length 11 ±2.2 19 ± 3.0 21 ± 5.9 20 ± 2.3
Breadth 1.4±0.5 2.1±0.5 1.4±0.8 1.4±0.5
Seed (mm)
Length 5.6±0.7 5.9±0.9 4.3±0.4 6.3±1.1
Breadth 1.9±0.5 1.9±0.4 1.5±0.7 1.7±0.5
Caruncle 3.1±0.8 3.0±0.7 2.2±0.5 3.5±0.7
Fig. 1. Polygala sekhukhuniensis. (A) Flowering plant, showing habit of above-ground parts; (B) Mature leaf; (C) Flower, lateral view; (D) Flower, lateral view with
wing-like sepal removed; (E) Old flower, lateral view with wing-like sepal removed to show capsule; (F) Ovary with style and stigma; (G) Androecium, showing
staminal tube joined at base to two upper petals; (H) Seed. Voucher: Van Wyk 13031. Scale bar 10 mm (A), 2 mm (BE), or 1 mm (FH). Artist: Daleen Roodt.
347S.J. Siebert et al. / South African Journal of Botany 76 (2010) 345353
6a. Stems greyish; carina 3 mm long with crest 1 mm long P. seminuda
6b. Stems green; carina 6 mm long with crest up to 3 mm long 7
7a. Racemes longer than 100 mm; pedicels 23 mm long P. hottentotta
7b. Racemes shorter than 100 mm; pedicels 12 mm long 8
8a. Flowers crowded in terminal clusters on raceme P. gracilenta
8b. Flowers evenly spaced along entire raceme 9
9a. Leaves 1622× 1.53.0 mm, apex acute; flowers with crest
white, pink or purple, alae whitish to yellowish with dark
green veins
P. leptophylla
9b. Leaves 915 × 1.02.5 mm, apex obtuse or retuse; flowers
with crest purple, alae pale pink with dark pink veins
P.
sekhukhuniensis
3.2. Palynology
In a review of Polygala palynological literature (Banks et al.,
2008), no mention is made of descriptions, drawings, or
micrographs of pollen belonging to the species compared in this
paper. Hence, the SEM observations on pollen grains presented
here are the first for the four taxa concerned (Fig. 3).
Pollen of P. sekhukhuniensis is isopolar and spheroidal in
equatorial view; P×E=(23)25.64(27)× (24)25.06(26)
μm; P/E =1.02 (Table 2). Apertures 22, zonocolpor ate.
Fig. 2. Stereomicroscope micrographs of seed and caruncle. (A) Polygala sekhukhuniensis, membranous appendage (me) 2.9 mm and chitinous appendage (ch) 0.8 mm,
hairs cover seed surface completely; (B) P. leptophylla var. leptophylla, me 2.7 mm and ch 1.6 mm, hairs cover two-thirds of the seed surface; (C) P. hottentotta, me 2.1 mm
and ch 1.0 mm, hairs cover seed surface completely; (D) P. seminuda, me 3.8 mm and ch 2.5 mm, hairs cover half of the seed surface. Scale bars 1 mm.
348 S.J. Siebert et al. / South African Journal of Botany 76 (2010) 345353
Ectoapertures (17)18.6(21) × (1)1.2(3) μm; endoapertures
(1)4.8(6) μm × endocingulate. Surface ornam entation psilate
to finely granular, with scattered lumina in apocolpial areas that
are small (less than 1 μm). Opercula present.
Pollen grains of P. sekhukhuniensis,andP. hottentotta differ
from P. leptophylla var. leptophylla and P. seminuda in shape,
namely oblate versus prolate and prolate-spheroidal respective-
ly (Fig. 3). Opercula over the endoapertures of P. sekhukhu-
niensis, P. hottentotta and P. seminuda are pronounced, mostly
so in the new species (Fig. 4). Although the shape of pollen
grains of P. sekhukhuniensis and P. hottentotta are very similar,
the former has smaller grains (Table 2).
Fig. 3. SEM micrographs of pollen. Polar view to the left. (A) Polygala sekhukhuniensis; (B) P. leptophylla var. leptophylla; (C) P. hottentotta; (D) P. seminuda.
Scale bars 10 μm.
349S.J. Siebert et al. / South African Journal of Botany 76 (2010) 345353
3.3. Distribution and ecology
Polygala sekhukhuniensis is only known from Sekhukhune-
land (Fig. 4). P. sekhukhuniensis and P. hottentotta occur on
patches of anomalous soils in this region sparsely vegetated
soils that are mineralized and rich in heavy metals (Table 3).
These soils are clayey and highly erodible, in contrast to the
rocky, sandy soil on mountain slopes that is preferred by P.
leptophylla var. leptophylla. These anomalies are akin to erosion
gulleys, but in this case not due to human disturbance, but caused
by natural erosion processes due to a weak soil structure and
associated high water dispersability (Mason, 1959). According to
Siebert et al. (2002) and Mandiwana et al. (2007), plant
communities typical of these eroded sites in Sekhukhuneland
include heavy metal accumulating or excluding species (Table 4).
Polygala sekhukhuniensis is such a species, specifically
adapted to grow in erosion gulleys of exposed heavy metal
contaminated (ultramafic) soil. Hence, the opinion that the
ecological species concept is an essential part of the biological
species concept (Grant, 1992) is supported here. Asymmetry and
within-plant variance are higher between specimens of the same
species in the contact zone between ultramafic and normal soils
(Alados et al., 1999). This variance provides genetic material for
natural selection and subsequent reproducti ve isolation.
In the case of P. sekhukhuniensis, an open niche with an
anomalous Ca-rich substrate (14.68% =146 800 ppm) in an
otherwise typical environment of soils rich in Mg (8.48% =
Table 2
Mean measurements (μm)/counts of selected pollen characters of Polygala sekhukhuniensis and three closely related species (n=10/species).
Polygala sekhukhuniensis Polygala leptophylla Polygala hottentotta Polygala seminuda
Nr. of apertures 22 20 21 21
Polar length 25.64 ±1.3 40.54±2.9 30.29±1.2 33.34±3.9
Equatorial breadth 25.06±0.9 25.98±1.8 33.27±1.4 29.66±1.5
P/E 1.02 1.56 0.91 1.12
Polar diameter 14.79 ±1.3 18.25±3.7 16.84±1.3 17.93±1.5
Ectoapertures lxw 18.6 × 1.2 30.6×0.8 22.9×1.1 26.4 ×1.3
Endoapertures lxw 4.8 ×endocingulate 2.1×endocingulate 6.9×endocingulate 5.3×endocingulate
Fig. 4. Known distribution of Polygala sekhukhuniensis.
Table 3
Environmental variables unique to plant communities containing either
Polygala sekhukhuniensis or P. hottentotta in the Sekhukhuneland Centre of
Plant Endemism.
Characteristics P. sekhukhuniensis P. hottentotta
Slope (°) 4 (07) 8 (015)
Aspect N, S, E, W S, W
Rock cover (%) 30 (555) 45 (2070)
Rock size (mm) 150 (100500) 450 (3001500)
Soil type Valsrivier, Prieska,
Mayo, Bonheim
Glenrosa, Mispah,
Valsrivier
Geology Norite, pyroxenite,
gabbro, alluvium
Pyroxenite, magnetite
Topographical position Footslope, valley Footslope, crest
Vegetation type Tall, sparse shrubland Short, open shrubland
350 S.J. Siebert et al. / South African Journal of Botany 76 (2010) 345353
84 800 ppm) and poor in Ca (2.13% = 21 300 ppm) (Fig. 5A),
probably favoured speciation. Similar to limestone, the soils
inhabited by P. sekhukhuniensis have a high Ca:Mg ratio (2:1),
which d iffer significantly from the soil substrate of P.
hottentotta (1Ca:4 Mg) in the study area (Fig. 5A). As a
consequence P. sekhukh uniensis has a leaf Ca:Mg ratio of 16:1,
and P. hottentotta a ratio of 4:1 (Fig. 6A). O'Dell et al. (2006)
have shown that the ability to maintain high leaf Ca:Mg is a key
evolutionary change needed for survival on heavy metal soil
and represents the physiological feature distinguishing the plant
species adapted to heavy metal soils from their non-adapted
congeners.
Furthermore, soils in which the two Polygala taxa grow have
high concentrations of total Cr, Al and Fe (Fig. 5A,B), with
Polygala hottentotta holding higher concentrations of Al and Fe
in its roots, stem s and leaves , but not at levels that could be
considered as accumulating (Fig. 6B). P. sekhukhuniensis
excludes these metals more effectively, despite the associated
soil concentrations being higher (Fig. 5B). Overall it seem s that
P. sekhukhuniensis is the better excluder of heavy metals due to
higher leaf Ca concentrations as a previously confirmed pro-
tection and survival mechanism (Konstantinou and Babalonas,
1996).
It is suggested that P. sekhukhuniensis originated as an
ecotype of and has developed from its suggested closest
relative, P. leptophylla, which occurs on nearby mountain
slopes. This theory was already proposed for Sekhukhuneland
endemics by Knowles and Witkowski (2000), suggesting that
the genetic properties of indi viduals occurring on iron-rich
mountain slopes have the potential to immigrate to, adapt to and
colonize ultramafic soils rich in heavy metals. It is hypothesized
that P. sekhukhuniensis is a typical edaphic specialist which
may well have speciated recently, after the Pleistocene (Reeves
et al., 1983). It probably prefers the competition-free, open
Table 4
Prominent associated taxa recorded for plant communities containing either
Polygala sekhukhuniensis or P. hottentotta in the Sekhukhuneland Centre of
Plant Endemism (Siebert et al., 2002).
Life forms Species dominant in:
P. sekhukhuniensis community P. hottentotta community
Grass layer Aristida congesta Diheteropogon amplectens
Fingerhuthia africana Loudetia simplex
Stipagrostis hirtugluma Themeda triandra
Forb layer Aloe cryptopoda Berkheya insignis
Dicoma gerrardii Jamesbrittenia macrantha
Gnidia polycephala Petalidium oblongifolium
Shrub layer Euclea sekhukhuniensis Searsia sekhukhuniensis
Grewia vernicosa Tinnea rhodesiana
Searsia keetii Vitex obovata
Tree layer Acacia sp. nov. Bolusanthus speciosus
Euphorbia tirucalli Combretum hereroense
Fig. 5. Chemical analyses of five soil samples collected from the root zone (300 mm deep) for each of Polygala sekhukhuniensis and P. hottentotta. Bulked sample per
species.
351S.J. Siebert et al. / South African Journal of Botany 76 (2010) 345353
niches of eroded, toxic ultramafic soils where it has a
physiological advantage requiring high levels of Ca to tolerate
heavy metals (Fig. 6A). P. sekhukhuniensis,likeEuclea
sekhukhuniensis, can therefore be considered a heavy metal
excluding, calciotrophic species.
3.4. Conservation status
Polygala sekhukhuniensis has a small geographical range in
Sekhukhune Plains Bushveld (Siebert et al., 2002), and limited
area of occupancy within natural erosion gulleys (dongas). Most
of its habitat is under threat from slimes dams and rock dumps
associated with the min ing industry, as these eroded areas are
considered to be of low conservation status. P. sekhukh uniensis
is not formally protected in any conservation area. Populations
of the species should therefore be closely monitored and its Red
Data List assessment assessed. The conservation value of P.
sekhukhuniensis is considered relatively high, as it could
possibly be used as a keystone species in the ecology of mine
dumps due to its internal mechanism of excluding heavy metals.
3.5. Etymology, vernacular names and uses
The specific epithet refers to the geographical area the
species occurs in, namely Sekhukhuneland, a region named for
King Sekhukhune(-i) I (18141882) of the Bapedi tribe. For
consistency, and in line with existing Latin epithets of plants
named after the region, it is based on the alternative spelling,
Sekhukhunila nd. We would like to propose the names
sekhukhune milkwort and sekhukhunebloukappie as Eng-
lish and Afrikaans vernacular names, respectively. The
recorded Northern-Sotho names for the plant are mabošêkgo
and mogaletsaoi, which differs from the vernacular name, pe-
loterri, given to the closely related P. leptophylla var.
leptophylla.
Burnt ashes are mixed with snuff for flavouring (Barnard
209) and the roots are cooked and given to forgetful people to
make the mind sensible (Barnard 135).
3.6. Additional specimens examined
LIMPOPO. 2429 (Zebediela): Mankopanie, Farm Hoera-
roep (BD), Barnard 209 (PRE); Bewaarkloof, Potlake Nature
Reserve (BD), Van Rooyen 2339 (PRE); Potlake Nature Reserve
(BD), Matthée 1044 (PRU); Farm Geeneinde, Sekukuniland (
DB), Barnard & Mogg 737 (PRE); Sekhukhuniland, Farm Parys
(DB), Barnard & Mogg 738 (PRE).
MPUMALANGA. 2430 (Pilgrim's Rest): Maandagshoek
(CA), Kritzinger 118 (PRE, PRU); Steelpoort, Eastern Chrome
Mines, valley beneath mountain in the Winterveld mine area
Fig. 6. Chemical analyses of plant material from five individuals each of Polygala sekhukhuniensis and P. hottentotta (young and old growth). Bulked sample per
species.
352 S.J. Siebert et al. / South African Journal of Botany 76 (2010) 345353
(CA), Siebert 44 9 (PRU); Sekukuni (CC), Barnard 135
(PRE); Dwarsrivier, 5 km on turn-off to Lydenbu rg from
Stofberg-Steelpoort road (CC), Jordaan 782 (PRE); 2 km
from Spitskop turn-off south on Steelpoort-Lydenburg road (
CC), Burgoyne 6015 (PRE); Thornecliffe Chrome Mine, hill
west of office ( CC), Van Wyk & Siebert 12982 (PRU);
Steelpoort, Ferrochrome Holdings, 4 km to east of Spitskop on
waytoRoossenekal(CC), Siebert 337 (PRU); Turnoff to
Thornecliff e Mine from Lydenbur g, Steelp oort road ( CC),
Van Wyk & Siebert 13311 (PRU); Road between Steelpoort
and Kennedy's Vale (CC), Siebert & Van Wyk 1379 (PRU).
Acknowledgements
The autho rs are indebted to Ma ggi Loubser, Geology
Department, University of Pretoria, for assistance with XRF
analyses and to Nina van Vliet, Department of Soil, Climate and
Water, Pretoria, for assistance with AAS and ICP-MS analyses.
We would like to thank Hugh Glen, for translating the
diagnoses into Latin, Kevin Balkwill for constructive comments
on a draft of the manuscript, Hester Steyn, for preparing the
distribution map and Daleen Roodt for the line drawings. The
National Research Foundation, University of Pretoria and South
African National Biodiversity Institute provided financial
support.
References
Alados, C.L., Navarro, T., Cabezudo, B., 1999. Tolerance assessment of Cistus
ladanifer to serpentine soils by developmental stability analysis. Plant
Ecology 143, 5166.
Angiosperm Phylogeny Group (APG) II, 2003. An update of the Angiosperm
Phylogeny Group classification for the orders and families of flowering
plants: APG II. Botanical Journal of the Linnean Society 141, 399436.
Balkwill, K., Campbell-Young, G.J., 2001. Taxonomic s tudies in the
Acanthaceae: Peristrophe serpenticola a new species from the Great
Dyke, Zimbabwe. South African Journal of Science 97, 551554.
Banks, H., Klitgaard, B.B., Claxton, F., Forest, F., Crane, P.R., 2008. Pollen
morphology of the family Polygalaceae (Fabales). Botanical Journal of the
Linnean Society 156, 253289.
Bredenkamp, C.L., 2000. Polygalaceae (RosidaePolygalales). In: Leistner, O.A.
(Ed.), Seed plants of southern Africa: families and genera. : Strelitzia, vol. 10.
National Botanical Institute, Pretoria, p. 450.
Burrows, S.M., Burrows, J.E., 2008. Three new species of Asparagus
(Asparagaceae) from South Africa, with notes on other taxa. Bothalia 38,
2329.
Exell, A.W., 1960. Polygalaceae. Flora Zambesiaca 1, 303336.
Grant, V., 1992. Comments on the ecological species concept. Taxon 41,
310312.
Hankey, A., Buys, M.H., Lebatha, P.D., 2008. Drimiopsis linioseta, a new
species from the Sekhukhuneland Centre of Endemism, South Africa.
Bothalia 38, 7274.
Knowles, L., Witkowski, E.T.F., 2000. Conservation biology of the succulent
shrub, Euphorbia barnardii, a serpentine endemic of the Northern Province,
South Africa. Austral Ecology 25, 241252.
Konstantinou, M., Babalonas, D., 1996. Metal uptake by Caryophyllaceae
species from metalliferous soils in northern Greece. Plant Systematics and
Evolution 203, 110.
Mabberley, D.J., 1997. The plant- book. Cambri dge University Pre ss,
Cambridge.
Mandiwana, K.L., Panichev, N., Kataeva, M., Siebert, S.J., 2007. The solubility
of Cr(III) and Cr(VI) compounds in soil and their availability to plants.
Journal of Hazardous Materials 147, 540545.
Manning, J.C., Goldblatt, P., 2009. Three new species of Gladiolus (Iridaceae)
from South Africa, a major range extension for G. rubellus and taxonomic
notes for the genus in southern and tropical Africa. Bothalia 39, 3745.
Mason, R., 1959. Later Pleistocene stratigraphy in the Transvaal. South African
Archaeological Bulletin 14, 38.
Massoura, S.T., Echevarria, G., Leclerc-Cessac, E., Morel, J.L., 2004. Response
of excluder, indicator, and hyperaccumulator plants to nickel availability in
soils. Australian Journal of Soil Research 42, 933938.
Morrey, D.R., Balkwill, K., Balkwill, M.-J., 1989. Studies on serpentine flora:
preliminary analyses of soils and vegetation associated with serpentine rock
formations in the southeastern Transvaal. South African Journal of Botany
55, 171177.
O'Dell, R.E., James, J.J., Richards, J.H., 2006. Congeneric serpentine and
nonserpentine shrubs differ more in leaf Ca:Mg than in tolerance of low N,
low P, or heavy metals. Plant and Soil 280, 4964.
Paiva, J.A.R., 1998. Polygalarum Africanarum et Madagascariensium prodro-
mus atque gerontogaei generic Heterosamara Kuntze, a genere Polygala L.
segregate et a nobis denuo recepti, synopsis monographica. Fontqueria 50,
1346.
Prenner
, G., 2004. Floral development in Polygala myrtifolia (Polygalaceae)
and its similarities with Leguminosae. Plant Systematics and Evolution 249,
6776.
Punt, W., Blackmore, S., Nilsson, S., Le Thomas, A., 1994. Glossary of pollen
and spore terminology. LPP Foundation, Utrecht.
Reeves, R.D., Brooks, R.R., Dudley, T.R., 1983. Uptake of nickel by species of
Alyssum, Bornmuellera, and other genera of the old world tribus Alysseae.
Taxon 32, 184192.
Retief, E., Siebert, S.J., Van Wyk, A.E., 2008. A new species of Euclea
(Ebenaceae) from ultramafic soils in Sekhukhuneland, South Africa, with
notes on its ecology. Bothalia 38, 3137.
Siebert, S.J., Van Wyk, A.E., Bredenkamp, G.J., 2001. Endemism in the flora of
ultramafic areas of Sekhukhuneland, South Africa. South African Journal of
Science 97, 529532.
Siebert, S.J., Van Wyk, A.E., Bredenkamp, G.J., 2 002. The physical
environment and major vegetation units of the Sekhukhuneland Centre of
Plant Endemism, South Africa. South African Journal of Botany 68,
127142.
Van Wyk, A.E., Smith, G.F., 2001. Regions of floristic endemism in southern
Africa. Umdaus Press, Hatfield.
Venter, H.J.T., Winter, P.J.D., Verhoeven, R.L., Archer, R.H., 2007. Raphio-
nacme villicorona (Apocynaceae: Periplocoideae), a new species from the
Sekhukhuneland Centre of Plant Endemism, South Africa. South African
Journal of Botany 73, 97101.
Vosa, C.G., 2007. Prototulbaghia, a new genus of the Alliaceae family from the
Leolo Mountains in Sekhukhuneland, South Africa. Caryologia 60,
273278.
Wild, H., 1965. The flora of the Great Dyke of Southern Rhodesia with special
reference to the serpentine soils. Kirkia 5, 4986.
Williamson, S.D., Robinson, E.R., Balkwill, K., 1997. Evolution of two
serpentine endemic taxa in Mpumalanga. South African Journal of Botany
63, 507513.
Yang, X.-E., Jin, X.-F., Feng, Y., Islam, E., 2005. Molecular mechanisms and
genetic basis of heavy metal tolerance/hyperaccumulation in plants. Journal
of Integrative Plant Biology 47, 10251035.
Edited by B-E Van Wyk
353S.J. Siebert et al. / South African Journal of Botany 76 (2010) 345353
... Consequently, through the process of speciation and natural selection, carbonate soils harbour unique plant communities with variable levels of endemism (Kruckeberg, 1969;Kruckeberg & Rabinowitz, 1985;Kruckeberg, 1986;Willis et al., 1996a;Cowling & Hilton-Taylor, 1997;Zhu et al., 2003;Qin et al., 2012;Smyčka et al., 2017). Endemic plant species on unusual soils are generally referred to as edaphic endemics, edaphic specialists or habitat specialists (Cowling & Holmes, 1992;Van Wyk et al., 2010;Magee et al., 2011;Goldblatt & Manning, 2012). Plants that are adapted to occupy Ca-rich habitats are referred to as calcicoles, calcicolous plants, calciphiles or calcicolous flora (Tansley, 1917;De Silva, 1934;Reinhardt et al., 2013;Rogers et al., 2018). ...
... Plant communities growing on carbonate soils are characterised by distinct species assemblages, high species richness and contribute significantly to regional as well as global plant diversity (Kruckeberg, 1969;Cowling, 1990;Zhu et al., 1998;Pärtel, 2002 Zietsman & Bredenkamp, 2007) and habitat loss (Willis et al., 1996a;Van Buren & Harper, 2003;Lu et al., 2016). Conservation of calcicolous floras remains challenging in Africa, due to restricted geographical plant distribution ranges and specific habitat preferences (Loehle, 2006;Retief et al., 2008;Van Wyk et al., 2010;Goldblatt & Manning, 2013). Thus, the second objective was to identify knowledge gaps and provide new perspectives by asking 86 significant questions that may contribute to future botanical research or guide conservation and management efforts of calcicolous plant communities in Africa. ...
... These included the provision of food and liquor to humans (Van Wyk et al., 2010), forage provision to livestock and game (Abd El-Ghani & Marei, 2007;Gamoun et al., 2010;Radloff et al., 2010;Gamoun, 2013;Ratovonamana et al., 2013) and the potential to assist with mine dump rehabilitation, especially those species that had the ability to exclude heavy metals (Retief et al., 2008). Some endemic species were found to have a medicinal value (Abd El-Ghani & Marei, 2007;Van Wyk et al., 2010). ...
Thesis
Full-text available
The Griqualand West Centre (GWC) of plant endemism harbours a unique flora of which 24 species are endemic. Heterogeneous geology, climate and topography are considered drivers of the unique flora and local endemism. However, these drivers have not yet been investigated and our understanding of the effects thereof on vegetation dynamics remains poor. Four mountain ecosystems, each underlain by different rock types and with distinct climatic patterns, provided a setting to investigate the effects of ecological drivers shaping vegetation dynamics of this semi-arid area. Therefore, the primary aim of this study was to disentangle the effects of rainfall and geology, through soil properties related to the underlying geological parent material, as drivers of floristic patterns, plant diversity and structure, biomass production, and the relationship between diversity and biomass production. The objectives of this study were to (i) redefine the borders of GWC to establish which mountain ranges fall within the centre by using a MaxEnt spatial model based on geology, climate and topography in combination with distribution data of GWC endemics, (ii) describe the flora within the newly redefined borders of GWC based on dominant plant families and -species, indicator plant species, endemic species and species composition, (iii) compare soil properties, rainfall, plant diversity and structure between mountain ecosystems to test whether mountains, within the newly defined borders of GWC, differ significantly from each other, (iv) determine whether soil properties, rainfall or a combination thereof act as drivers of plant diversity and structural differences between mountains, (v) test for differences in total biomass production (above ground green plant material and debris), live biomass production (only live green above ground plant material) and respective plant functional group (PFG) biomass production between the four mountain rangelands, (vi) relate differences to specific soil properties and rainfall to identify the strongest drivers of biomass production, (vii) investigate diversity-biomass relationships for total plant species and for species representing different PFGs, and (viii) present an optimal range of biomass production at which herbaceous species diversity can be maintained at regional scale. Results obtained from this study revealed that each mountain plant community was characterised by unique herbaceous plant communities with specific indicator plant species, driven by soil properties and rainfall. Herbaceous plant composition, density, height, cover and shrub frequencies were related to a combination of soil properties and mean annual rainfall. However, plant diversity, and grass, lignified forb and tree frequencies, as well as woody plant height and canopy area, could only be related to soil properties. Grasses, lignified forbs and herbaceous forbs contributed to biomass production in descending order. At regional and local scales, diversity-productivity relationships followed non-linear trends. However, optimum biomass production was reached at highest diversity. Semi-arid mountain landscapes in GWC provide important ecosystem services through their unique plant diversity. It is necessary to follow a holistic, multi-disciplinary conservation and management approach to not only manage for species diversity, but to conserve the underlying environmental drivers in semi-arid mountain plant communities.
... He also developed the first database of the collection which is now hosted on the BRAHMS platform. Regional geoecological studies have led to the discovery and description of rare and endemic edaphic specialists from Sekhukhuneland in Limpopo (Siebert et al. 2010). A phylogenetic study was also undertaken of the southern African Andropogoneae (Poaceae) to determine the potential of gene flow from Saccharum hybrids to wild relatives (Snyman et al. 2018 Taxonomists at NWU have made a concerted effort to contribute to national (Siebert et al. 2010) and regional (Struwig et al. 2015) efforts to categorize and name plants. ...
... Regional geoecological studies have led to the discovery and description of rare and endemic edaphic specialists from Sekhukhuneland in Limpopo (Siebert et al. 2010). A phylogenetic study was also undertaken of the southern African Andropogoneae (Poaceae) to determine the potential of gene flow from Saccharum hybrids to wild relatives (Snyman et al. 2018 Taxonomists at NWU have made a concerted effort to contribute to national (Siebert et al. 2010) and regional (Struwig et al. 2015) efforts to categorize and name plants. Subsequently, in recent years, closer collaboration has been sought and established with the Pretoria National Herbarium of the South African National Biodiversity Institute (PRE) through the appointment of two of their taxonomic experts as extraordinary lectures to improve the functioning of the herbarium, provide database support and stimulate collaborative fieldwork. ...
Conference Paper
Full-text available
The A.P. Goossens Herbarium (PUC) was founded by Antonie Goossens in 1932 and today it holds over 30 000 specimens from central South Africa. A brief history of herbarium establishment, development as well as its educational purposes, results of scientific studies (taxonomical, ecological and biogeographical) and current status and problems are described and discussed.
... Regional geoecological studies have led to the discovery and description of rare and endemic edaphic specialists from Sekhukhuneland in Limpopo (Siebert et al. 2010). A phylogenetic study was also undertaken of the southern African Andropogoneae (Poaceae) to determine the potential of gene flow from Saccharum hybrids to wild relatives (Snyman et al. 2018 Taxonomists at NWU have made a concerted effort to contribute to national (Siebert et al. 2010) and regional (Struwig et al. 2015) ...
... Regional geoecological studies have led to the discovery and description of rare and endemic edaphic specialists from Sekhukhuneland in Limpopo (Siebert et al. 2010). A phylogenetic study was also undertaken of the southern African Andropogoneae (Poaceae) to determine the potential of gene flow from Saccharum hybrids to wild relatives (Snyman et al. 2018 Taxonomists at NWU have made a concerted effort to contribute to national (Siebert et al. 2010) and regional (Struwig et al. 2015) ...
... The newly collected specimens were deposited in Guilan (GUH) (Holotype) and Yasouj (Isotype) Universities Herbaria. The new species was identified using different Floras and related articles (Boissier 1867: 76, Shishkin 1964: 118, Cullen 1965: 533, Chrtek & Krisa 1977, Peşmen 1980, Davis et al. 1988, Jalilian 2005, Eren et al. 2008, Raabe et al. 2009, Siebert et al. 2010 Wahlert et al. 2017, Dönmez & Uðurlu Aydin 2018, Sarvi et al. 2020. It was also compared with other species of Polygala in central herbarium of Tehran University (TUH), herbarium of Shiraz University (HSHU), Iranian research institute of plant protection (IRAN), herbarium of research institute of forests and rangelands (TARI) and herbarium of animal & natural resources research center of Hormozgan province. ...
Article
A new species, Polygala guilanica (Polygalaceae), is described from Kooh-Roubar, Gasht-Roodkhan protected area in W Guilan province, N Iran. The new species is distinguished based on its prostrate stem; small, lanceolate, ovate and rhombic, sessile leaves; tiny pink-white flowers in lax terminal racemes; 6 or 8 stamens with sessile anthers, a filiform style and brownish seed with 3-lobed caruncle. Detailed morphological description of the new species, photographs and distribution map are provided. Polygala guilanica is compared with the morphologically closest species: P. kurdica, P. hohenackeriana and P. anatolica. An updated identification key is provided for the Iranian species.
... The newly collected specimens were deposited in Guilan (GUH) (Holotype) and Yasouj (Isotype) Universities Herbaria. The new species was identified using different Floras and related articles (Boissier 1867: 76, Shishkin 1964: 118, Cullen 1965: 533, Chrtek & Krisa 1977, Peşmen 1980, Davis et al. 1988, Jalilian 2005, Eren et al. 2008, Raabe et al. 2009, Siebert et al. 2010 Wahlert et al. 2017, Dönmez & Uðurlu Aydin 2018, Sarvi et al. 2020. It was also compared with other species of Polygala in central herbarium of Tehran University (TUH), herbarium of Shiraz University (HSHU), Iranian research institute of plant protection (IRAN), herbarium of research institute of forests and rangelands (TARI) and herbarium of animal & natural resources research center of Hormozgan province. ...
Article
Full-text available
A new species, Polygala guilanica (Polygalaceae), is described from Kooh-Roubar, Gasht-Roodkhan protected area in W Guilan province, N Iran. The new species is distinguished based on its prostrate stem; small, lanceolate, ovate and rhombic, sessile leaves; tiny pink-white flowers in lax terminal racemes; 6 or 8 stamens with sessile anthers, a filiform style and brownish seed with 3-lobed caruncle. Detailed morphological description of the new species, photographs and distribution map are provided. Polygala guilanica is compared with the morphologically closest species: P. kurdica, P. hohenackeriana and P. anatolica. An updated identification key is provided for the Iranian species.
... Indicator plant species (Table 7) are characterized by high relative frequency of occurrence in a specific mountain habitat (specificity) and thus were primarily found in that habitat in high numbers (Dufrêne and Legendre 1997). In contrast to common species, indicator plant species provide valuable ecological information on various species groups of different plant communities (Dufrêne and Legendre 1997), especially with respect to their habitat preferences and adaptations to persist under certain environmental conditions (Siebert et al. 2010). ...
Article
Full-text available
Van Staden N, Siebert SJ, Cilliers DP, Wilsenach D, Frisby AW. 2020. Floristic analysis of semi-arid mountain ecosystems of the Griqualand West centre of plant endemism, Northern Cape, South Africa. Biodiversitas 21: 1989-2002. The Griqualand West Centre (GWC) is one of 13 centres of plant endemism in South Africa. Despite its unique flora, it remains poorly conserved and studied. A recent study identified an extensive geographical core area for the GWC, but endemic plant species were found to be absent from certain parts within these borders. To address this, we refined the current GWC borders based on an ecological niche model, which predicted that endemic species are restricted to four mountain ranges within GWC. Mountain floras within these refined borders were then floristically compared to assess whether they are hotspots of endemicity. Floristically, the Asteraceae, Fabaceae, Malvaceae, and Poaceae were the dominant plant families. Mountain ecosystems differed from one another at species level, with indicator species explaining the compositional differences. Distribution patterns of indicator species were determined by mean annual precipitation, Ca: Mg ratios, soil pH, cation exchange capacity, iron, and sand content. These environmental factors are possible drivers of niche partitioning, environmental filtering and habitat specialization in each mountain ecosystem. Limestone and banded ironstone habitats were identified as conservation priority areas, since they contained the highest numbers of rare and threatened GWC restricted-range species, of which six were narrow endemics.
... Boloponera ikemkha was found within the Sekhukhuneland Centre of Plant Endemism (SCPE), an area recognised for its unique plant diversity (Siebert et al. 2002(Siebert et al. , 2010 but as yet very poorly known from an invertebrate perspective. The type locality of B. ikemkha is approximately 3400 km SSE of that of B. vicans (see Figure 5) and in a markedly different habitat with very different climatic conditions. ...
Article
Full-text available
During an environmental impact assessment survey of a proposed tailings storage facility for a platinum mine in Sekhukhuneland, South Africa, five adult and five larval specimens of a new species of Boloponera were found while excavating soil to a depth of 10–15 cm at the base of a tree in riparian woodland. These specimens represent a 3400 km range extension and the first reported record of the genus since its description in 2006, which was based on a single specimen collected in the Central African Republic in 2001. A description of the worker and ergatoid queen of Boloponeraikemkhasp. n . is presented, with a description of the mature larva and a key to distinguish workers of the two currently known species of the genus. The taxonomic relationships of Boloponera are discussed with respect to several confirmed and newly identified autapomorphies that support its retention as a distinct genus, although closely related to Plectroctena and Loboponera . A preliminary assessment of the conservation status and discussion of potential threats to the survival of B.ikemkha is also provided. Evaluation of current data under the IUCN Red List criteria would result in B.ikemkha being assessed as Critically Endangered, but further investigation is required to test the validity of placing it in this category.
Article
Full-text available
In this paper we celebrate this milestone by giving an overview of the history of the Department at this campus, including its recent establishment and expansion on the NWU Mahikeng Campus (formerly the University of NorthWest). A brief overview is presented of the advances in teaching and research over the years, and the development and relevance of the important plant collections in the botanical garden, two herbaria and the national diatom collection. The main emphasis of this contribution is, however, a reflection on the advance ment and significance of research conducted by various disciplines on plant and algae function, diversity and ecological restoration over the years. The different disciplines in Botany at NWU, from the oldest to the more recent, are Plant Taxonomy, Plant Ecophysiology, Terrestrial Plant Ecology, Aquatic Sci ences, Urban and Settlement Ecology, Geoecology, and Proteomics. Different aspects contributing to changes occurring in the environment, such as pollution, land degradation, urbanisation, overexploitation of resources and the subsequent effect of these on plant diversity and function are especially ad dressed in our current research. The results of our research inter alia led to solu tions for problems occurring in the landscape and contribute to the wellbeing of the people using the land and water by restoring important ecosystem services.
Thesis
EXTENDED THESIS ABSTRACT Context Understanding spatial patterns of biodiversity and species’ distributions is important for scientific theory, and for conservation and management of the natural world. Climatic variables are widely recognised as strong correlates of species richness over large spatial extents. Correlates of species richness at smaller extents (regional and landscape scales) are less well established, but environmental heterogeneity is widely thought to be important. A large number of environmental heterogeneity measures have been used, but in particular there is a growing interest in ‘geodiversity’, which I define here as the diversity of abiotic terrestrial and hydrological nature, comprising earth surface materials and landforms. Recent research has emphasised both geodiversity’s inherent value and its potential as a correlate and predictor of spatial biodiversity and species’ distribution patterns. However, despite this clear potential of geodiversity for improving our understanding of how patterns of life relate to environmental heterogeneity, its incorporation into biodiversity and species’ distribution modelling is substantially underdeveloped. In this thesis, using a macroecological approach I begin to address some of these knowledge gaps by analysing the relationships between geodiversity data, and its constituent ‘geofeatures’, and species richness and distributions for multiple taxa and across several scales (grain size and extent) and geographic locations. My main aims in this thesis are to more fully evaluate geodiversity itself, and improve our understanding of its role with respect to the spatial patterning of biodiversity, both conceptually and empirically. Locations and Spatial Scales Analyses were carried out within and across Great Britain (England, Scotland, and Wales) and Finland. The order of the four quantitative papers generally reflects the largest spatial extent (i.e. size of the study area) at which they were conducted, from national (PAPERS II and III) through landscape (PAPER IV), to the local scale (vegetation plots within a small upland river catchment; PAPER V). PAPER II is a study across several spatial extents (from landscape to national) and uses two grain sizes (1 km2 and 100 km2). PAPER I is a review paper that considers multiple scales and geographic locations conceptually. Time period Present day: data were from between 1995 and 2016 across all of the quantitative studies. Taxa Multiple: alien and native vascular plants across Great Britain (PAPER II); threatened bryophytes, beetles, fungi, lepidoptera, lichens, mammals, molluscs, and vascular plants across Finland (PAPER III); common and rare vascular plants across the Cairngorms, Scotland (PAPER IV); angiosperms, conifers, fungi, lichens, liverworts, lycophytes, mosses, and pteridophytes (and productivity) across an upland river catchment within the Cairngorms (PAPER V); and conceptual consideration of multiple taxa (PAPER I). Methods For studies in Great Britain, plant data were provided by the Botanical Society of Britain and Ireland (BSBI) for PAPERS II and IV, and by the Centre for Ecology and Hydrology (CEH) for PAPER V. The threatened species data in Finland were from Finnish Environment Institute (PAPER II). Species richness (PAPERS II, III, and IV), rarity-weighted richness (RWR; PAPER III), species’ distributions (PAPERS IV and V), and productivity (measured using NDVI from colour infrared aerial imagery; PAPER V) were all analysed using Boosted Regression Tree (BRT) modelling, allowing comparisons between studies. For geodiversity data in the British studies, I compiled geodiversity data on landforms, soils, hydrological and geological features using existing national datasets (e.g. British Geological Survey), and used a geomorphometric method to extract landform coverage data (landforms included: hollows, ridges, valleys, and peaks). These data were analysed alongside environmental data, which varied between papers, relating to climate, standard topography (e.g. slope; elevation), land use, and human population. The sources of other geodiversity data in Finland, and environmental data on topography and climate, came from a variety of sources, which are detailed within each paper. Results Geodiversity improved biodiversity and species’ distribution models throughout all of the quantitative analyses and generally declined in importance as spatial scale coarsened beyond the landscape scale. At most spatial scales and in most places, the roles of climate and/or coarse topography dominated, and geodiversity played a relatively small role, as was expected. Geodiversity, however, made consistent positive contributions to the models independently of traditionally used topographic metrics such as standard deviation of elevation and slope. Taxonomically, geodiversity: (i) was slightly more relevant for native vascular plants than alien in Great Britain (PAPER II); (ii) of similar relevance to common and rare vascular plants in the Scottish Highlands, except that the coverage of soil parent material was especially important for rare species’ distributions (PAPER IV); of similar relevance to most sessile taxa (angiosperms, fungi, mosses, liverworts, lichens, pteridophytes, and lycophytes; conifers were not related to geodiversity) in an upland Scottish river catchment (PAPER V); and more important for threatened vascular plants and bryophytes over other studied taxa in Finland (PAPER II). Geodiversity also improved models of productivity, and the variability in productivity, in PAPER V. Main conclusions and Future Directions Geodiversity improves our understanding of, and ability to model, the relationship between biodiversity and environmental heterogeneity at multiple spatial scales, by allowing us to get closer to the real-world conditions and processes that affect life. I found that the greatest benefit comes from measuring ‘geofeatures’, which describe the constituent parts of geodiversity separately, rather than as one combined variable. Automatically extracted landform data, the use of which is novel in ecology, biogeography and macroecology, proved particularly valuable throughout this body of work, and as too did data from expert geological and hydrological maps. The idea of ‘Conserving Nature’s Stage’ (CNS), and identifying areas that are most capable of supporting high biodiversity into the future, the benefits and caveats of which are discussed in this thesis, has recently emerged. It requires a sound empirical and conceptual basis, to which my research contributes. In this thesis, I have gone some way towards demonstrating the conceptual and empirical value of incorporating geodiversity into ecological analyses across multiple spatial scales, paving the way for this recent approach to be more extensively used for theoretical and applied purposes. I accomplished this by carrying out an assessment of existing geodiversity literature and, importantly, looking forwards to consider the prospects of geodiversity within ecology (PAPER I), supported by four quantitative studies. The conservation significance is emphasised in PAPER III. Much remains to be done, however, and future research directions are detailed in PAPER I. We need to develop predictive models to test the role of geodiversity across an array of geographical and taxonomic domains, as well as to assess metrics beyond species richness and species’ distributions. One example may involve beta diversity: does spatial turnover in species relate to spatial turnover in geofeatures? Fully analysing the role of geodiversity through time will also be important, including in relation to refugia, given predicted environmental changes in climate. In progressing with this line of enquiry, we will improve our knowledge and understanding of patterns of life on Earth and, specifically, how the geophysical landscape helps shape them.
Article
The first record of Polygala westii (Polygalaceae) from the North-West Province (South Africa) is reported on here. The species is very rare and has previously only been collected in two separate gatherings. This species is characterised by having bracts and bracteoles caducous, the anterior sepals connate at least as much as half of their length and seeds pubescent and carunculate. It grows in a habitat that are severely impacted by agriculture and mining. A preliminary assessment of its conservation status is given. A key to the South African species in the subsection (Tetrasepalae) to which P. westii belong is provided. The aim of this paper is to highlight the species in order to stimulate the lookout for more material of this species.
Article
Full-text available
Three new species of Gladiolus L. are described from South Africa. G. dolichosiphon is the second known member of series Blandus from the mountains of the Little Karoo in Western Cape, and is distinguished from other members of the long-tubed, pink-flowered G. carneus complex by its 5 or 6 linear leaves, creamy pink to salmon flowers with a tube 30-50 mm long and longer than the dorsal tepal, and its late summer flowering. G. karooicus from the Klein Roggeveld and the northern foothills of the Witteberg, is a spring-flowering species allied to G. permeabilis but has bright, canary-yellow flowers with the lower part of the lower tepals involute and conspicuously auriculate. G. reginae is an edaphic endemic of the Sekhuk-huneland Centre of Floristic Endemism in Mpumalanga, and flowers in autumn. It is evidently a glabrous member of section Densiflorus series Scabridus, distinguished by its long-tubed flowers, streaked with red on the lower tepals and blotched with red in the throat. Anomalously, however, it has the tubular inner bracts and large capsules diagnostic of section Ophiolyza series Oppositiflorus. A re-examination of the morphology suggests that series Scabridus is better placed in section Ophiolyza and a slightly revised classification of Gladiolus in southern Africa is proposed. We also propose the replacement name G. sulculatus for the Tanzanian species, G. sulcalus Goldblatt, a later homonym of G. sulcatus Lam. Finally, a recent sighting of what appears to be G. rubellus from northern Namibia constitutes the first record of this species in the country and a major range extension from its previous known occurrence in southeastern Botswana.
Article
Full-text available
DRIMIOPSIS LINIOSETA, A NEW SPECIES FROM THE SEKUKHUNELAND CENTRE OF ENDEMISM, SOUTH AFRICA
Article
Two taxa endemic to serpentine soils in Mpumalanga, Berkheya rehmannii Thell. var. rogersiana Thell. and B. coddii Rossl. (Asteraceae) were selected to investigate the mode of evolution of serpentine endemics using enzyme electrophoresis. Morphological studies based on herbarium material suggested that the serpentine endemic B. rehmannii var. rogersiana is probably closely related to B. rehmannii Thell. var. rehmannii which does not grow on serpentine. The electrophoretic investigation revealed high genetic identities between all taxa studied. However, the presence of unique alleles in var. rogersiana and large differences in allele frequencies between var. rogersiana and var. rehmannii suggest that these taxa are genetically distinct, that there is little or no gene flow between them and that they could be a progenitor-derivative pair. These data and the distribution data suggested that var. rogersiana is probably not neo-endemic nor palaeo-endemic but somewhere in between and B. coddii is probably palaeo-endemic and that they arose during independent evolutionary events. Thus, palaeo-endemism probably exists in the elements of the serpentine flora in Mpumalanga that were studied and both serpentine-tolerance and nickel accumulation have evolved more than once.
Article
In 1974, Wild collected a species of Peristrophe Nees from Vanad Pass on the Great Dyke of Zimbabwe. It does not match any of the known species and differs from the closely related P. decorticans K. Balkwill in leaf colour, size and colour of the flowers, texture of the hairs on the stems, size of the tertiary bracts, corolla tube, corolla lips and fruit. The new species, P. serpenticola K. Balkwill & G. J. Campbell-Young, is also similar to P. cernua Nees, but differs in size of the secondary bracts, corolla tube, corolla lips, fruits and seeds. P. serpenticola also differs from P. decorticans and P. cernua in tubercles and pattern of the testa surface. P. serpenticola is known only from serpentine soils of the Great Dyke and is thus considered a serpentine endemic. This species grows almost exclusively on termite mounds, which protects it from fire but does not seem to ameliorate the toxic nature of the soil. The IUCN conservation status of P. serpenticola is considered endangered, since the species exists in very small, localized populations, as is shown by the limited herbarium collections. The serpentine habitat on the Great Dyke is also under threat from mining.
Article
Soils sampled from sites adjacent to two disused mines in the Barberton district were found to be derived from ultrabasic rocks. The soils contained elevated concentrations of Ni and Cr. Although potentially phytotoxic, the soils supported a relatively diverse flora. Samples of six species collected from both sites had unusually high root and leaf tissue concentrations of Ni, suggesting physiological tolerance of the metal. Berkheya coddii Roessl. showed hyperaccumulation of Ni in leaves and is likely to be endemic to nickeliferous serpentine; it may also have value as a geobotanical indicator species. Only Sporobolus pectinatus Hack. and Sutera sp. aff. S. silenoides Hilliard absorbed Cr from soil solution and translocated it to leaves. Uptake and translocation of Cr is rare and the physiological mechanisms of Cr tolerance in these species remains unclear.