Genet Resour Crop Evol

https://doi.org/10.1007/s10722-020-00886-8 (0123456789().,-volV)
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RESEARCH ARTICLE

Diversity of grasses (Poaceae) in southern Africa,

with emphasis on the conservation of pasture genetic
resources
M. Trytsman . F. L. Müller . A. E. van Wyk

Received: 12 June 2019 / Accepted: 16 January 2020

Ó Springer Nature B.V. 2020

Abstract A renewed interest in the present state of
genebanks conserving pasture genetic resources
worldwide motivated this study to quantify the wealth
of grass (Poaceae) diversity indigenous to southern
Africa, here defined as South Africa, Lesotho and
Eswatini (previously Swaziland). Botanical occur-
rence records were extracted from BODATSA and
PHYTOBAS datasets to generate a list of grass species
indigenous to the study area. The phylogenetic
classification, growth form, photosynthetic pathway,
grazing status, endemism and conservational status
attributes were added to the 43,889 species level
records, sourced from published literature. Results
from the current study indicate that the subcontinent is
represented by eight subfamilies, 25 tribes, 151 genera
Electronic supplementary material The online version of
this article (https://doi.org/10.1007/s10722-020-00886-8) con-
tains supplementary material, which is available to authorized
users.
and 685 species, inferring that only 20% of the world’s
grass genera and 6% of world’s grass species are found
in the study area with Panicoideae the most speciose
subfamily. Paniceae is the only tribe with large
numbers of both C3 and C4 species and with several
species of high grazing value, therefore, was suggested
as a priority lineage in the collection and conservation
efforts of the South African National Forage Gene-
bank. This genebank conserves at present 73 genera
and 162 indigenous grass species, i.e. 48% and 24% of
the total number of taxa respectively, denoting the
current vulnerable status of grass genetic resources in
southern Africa. A need to therefore collect and
conserve grass genetic resources is emphasised, with
greater focus on the conservation of seed of well-
known pasture genera classified as endangered or
possibly extinct (mainly Panicum L. and Secale L.).
Keywords Endemism
Eswatini
Gramineae

Lesotho
South Africa

M. Trytsman (&)
F. L. Müller

Agricultural Research Council-Animal Production
Institute, Lynn East, PB X05, Pretoria 0039, South Africa Introduction
e-mail: mtrytsman@arc.agric.za

A. E. van Wyk Poaceae (Gramineae), the members of which are

Department of Plant and Soil Sciences, University of commonly referred to as grasses and bamboos, is
Pretoria, Pretoria 0002, South Africa considered as probably the most valuable plant family
A. E. van Wyk
to humankind (Bouchenak-Khelladi et al. 2010). The
National Herbarium, South African National Biodiversity family includes the economically important cereals,
Institute, Pretoria 0001, South Africa

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sugar crops, reeds, bamboos, forages and lawn grasses
(Hodkinson 2018). The success of this family can be
attributed to, amongst others, its adaptability to most
ecosystems, including arctic regions and at high
elevations uninhabitable by flowering plants (Tzvelev
1989), ecological dominance in many ecosystems, and
high species richness (Linder et al. 2018).
The important role of the Poaceae in sustainable
livestock production is well known, with several
genera housing important pasture species (Truter et al.
2015; Capstaff and Miller 2018). The exploration of
the potential of southern African grasses for pastures
began as early as the 1900s, with the past 50 years
described as a period where the function and value of
southern African grasses were studied by several
pasture researchers (Truter et al. 2015). An important
initiative towards the conservation of grass genetic
resources in southern Africa was a collection excur-
sion specially arranged for this purpose, to the Kruger
National Park in South Africa, during the early 1990s.
This ensured that selected ecotypes of important
pasture species such as Anthephora pubescens Nees,
Cenchrus ciliaris L., Chloris gayana Kunth, Cynodon
dactylon (L.) Pers., Digitaria eriantha Steud., Era-
grostis curvula (Schrad.) Nees and Panicum maximum
Jacq. were conserved in the South African National
Forage Genebank (SA-NFG) (Kruger et al. 1993).
However, similar to trends in the international
genebank community, especially those housing trop-
ical and sub-tropical plant genetic resources (Maass
and Pengelly 2019; Pengelly and Maass 2019), the
conservation of these plant genetic resources have
been under threat for the last 20 years. Generally, the
funding to manage and maintain forage genebanks
around the globe is on the decrease, resulting in many
important plant genetic resources potentially being
lost (Maass and Pengelly 2019). In the case of the SA-
NFG, the plant genetic resources maintained at the
facility was mainly threatened by the lack of funding
which resulted in unreliable storage and testing
facilities, coupled with a decline in trained personnel
capable of maintaining and evaluating the valuable
collection of plant genetic resources housed at the
facility. Also, the SA-NFG houses a large number and
diversity of plant genetic resources, many of which
currently are perceived to have minimal to no
agronomic potential due to the lack of information
regarding their pasture potential. These species were
collected with future breeding in mind.
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Genet Resour Crop Evol
The decline in the pasture breeding capacity in
South Africa (Truter et al. 2015) has resulted in many
of these species not being characterised and evaluated
for their pasture potential. Therefore, due to the lack of
information regarding their pasture potential these
species have not yet been included in breeding and
evaluation programs. These plant genetic resources
are, however, still important sources of genetic
material that could, under future bioclimatic condi-
tions become valuable resources for breeding of future
pasture species adapted to specific agro-ecological
conditions. As a result, these plant genetic resources
are maintained in perpetuity at the SA-NFG as a means
to maintain the plant genetic diversity that could
potentially be beneficial under future bioclimatic
conditions. Maintaining and conserving these plant
genetic resources however, puts significant financial
pressure on the SA-NFG. Recently, Pengelly and
Maass (2019) and Maass and Pengelly (2019) called
for the prioritization of germplasm as well as to
improve the efficiency in conserving current collec-
tions of plant genetic resources housed at genebanks
across the globe. This, in turn, is believed to reduce the
financial burdens on genebanks, by shifting efforts to
only priority species, and locally adapted varieties and
ecotypes within important pasture species with known
agronomic potential (Pengelly and Maass 2019; Maass
and Pengelly 2019). This will allow for the conserva-
tion of a larger genetic diversity within species with
known agronomic potential, which, in turn, will
benefit future breeding programs of these species.
An example of this can be found in domesticated
African grass species such as Sorghum bicolor (L.)
Moench and Pennisetum glaucum (L.) R.Br. that are
currently maintained and conserved for at least two-
thirds of their diversity of their wild relatives (Buckler
et al. 2001).
Van Wyk (1995) emphasized that the recognition
and interpretation of genetic variation in organisms is
at the heart of taxonomy and called on plant
taxonomists in Africa for the urgent naming of
infraspecific units. A striking example that highlighted
the importance of this appeal was the revision of
Agrostis eriantha Hack. var. planifolia Goossens &
Papendorf being reduced to synonymy under A. eri-
antha Hack. (Victor et al. 2012). A method to
prioritize taxonomic revision of South African plant
genera was developed by Victor et al. (2015) to reduce
the taxonomy-conservation disorder where indicators
Genet Resour Crop Evol
such as revision dates, insufficient data (including
taxonomic uncertainty) and endemism were proposed
as indicators. Furthermore, a literature review done in
the early 1990s showed that only a few key or
important indigenous grass species had complete
autecological studies with the exception of Themeda
triandra Forssk., Eragrostis curvula and Digitaria
eriantha (Shackleton 1991). A need for a coordinated,
systematic approach in basic ecological research of
grass species in different biomes was therefore
suggested.
The most recent taxonomic review of genera in
southern African grasses was within the temperate
genus Helictotrichon Besser s.l. (Mashau et al. 2010)
where two new species were identified. Germishuizen
and Meyer (2003) recorded major name revisions
within the genera Aristida L. [= Stipagrostis Nees],
Danthonia DC [= Merxmuellera Conert; = Kar-
roochloa Conert & Türpe; = Chaetobromus
Nees; = Centropodia Rchb.; = Dregeochloa Conert],
Phragmites Adans. [= Tribolium Desv.] and Rhynch-
elytrum Nees [= Melinis P.Beauv.] whilst Pentaschis-
tis (Nees) Spach and Setaria P.Beauv. were revised at
species level. However, Fish et al. (2015) indicated
that the genera Agrostis L., Anthoxanthum L., Cym-
bopogon Spreng., Cynodon Rich., Echinochloa
P.Beauv. and Puccinellia Parl. are also in need of
revision. Remarkably, a new species of Enneapogon
Desv. ex P.Beauv. was recently identified by Mashau
and Coetzee (2019) i.e. Enneapogon limpopoensis
Mashau, being possibly endemic to South Africa and
Zimbabwe.
The classification system followed in earlier con-
tributions on the identification of southern African
grasses (Gibbs Russell 1986; Ellis 1988; Gibbs Russell
et al. 1990) differ greatly from the present-day system
(Fish et al. 2015). Gibbs Russell et al. (1990)
recognised five subfamilies and 21 tribes whereas
Fish et al. (2015) recognized eight subfamilies and 24
tribes. Hence, one of the aims of the present study is to
provide fresh insights into southern African’s valuable
grass genetic resources within a modern classification
framework. For the present study, the most recent
worldwide phylogenetic classification of Soreng et al.
(2017) will be used for taxa found in the study area.
The identification manuals of southern African grasses
(Gibbs Russell et al. 1990; Fish et al. 2015) listed all
indigenous grasses and also described the photosyn-
thetic pathway and growth form, highlighting the
diversity of indigenous grasses at tribe and species
level. Linking attributes such as photosynthetic path-
way, growth form and grazing status with phyloge-
netic classification could assist in distinguishing taxa
with pasture potential. To ensure a similar outcome as
for the pasture potential appraisal of legumes (Legu-
minosae/Fabaceae) indigenous to South Africa,
Lesotho and Eswatini (previously Swaziland) (Tryts-
man et al. 2016, 2019), this paper takes stock of the
wealth of indigenous grass genetic resources with
added references to grass species currently used in
pasture systems. This, in turn, will help with efforts to
prioritise conservation of important grass genetic
resources at the SA-NFG, in line with the call by
Maass and Pengelly (2019) and Pengelly and Maass
(2019).
Methods
The Botanical Database of Southern Africa (BOD-
ATSA) maintained by the South African National
Biodiversity Institute’s (SANBI) and stored in the
BRAHMS platform (Le Roux et al. 2017) was
accessed on 2017/03/24 to extract the occurrence
records for Poaceae. The taxon and quarter degree grid
cell (QDGC) records were extracted and refined, i.e.
alien and naturalized species, species with no QDGC
reference, species present outside the study area,
namely South Africa, Lesotho and Eswatini, hence-
forth referred to as southern Africa [SA], invalid
botanical names, synonyms, as well as incomplete
taxa were removed from the dataset. Genus and
species names were verified using Fish et al. (2015) to
ensure that only species indigenous to SA were
included and naturalized or species recorded from
countries bordering the study area, were excluded. The
database does not reflect all herbarium records from
southern Africa, but mainly those housed in the
National Herbarium (PRE) in Pretoria and some of
its satellite herbaria, notably the KwaZulu-Natal
Herbarium (NH) in Durban and the Compton Herbar-
ium (NBG) in Cape Town. Despite its inherent
limitations, the meaningful results generated justify
the use of this database, the only one of its kind for the
study area (Trytsman et al. 2016).
In addition, the botanical survey records contained
in the PHYTOBAS database were accessed and
Poaceae records with GPS locations were extracted.
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 Genet Resour Crop Evol

Table 1 Botanical records of the Poaceae indigenous to South Africa, Lesotho and Eswatini contained in the Botanical Database of
southern Africa (BODATSA; maintained by SANBI) and PHYTOBAS (South African National Vegetation Data Archive) datasets
Database Date #QDGCs/GPS #Taxa #Records

BODATSA: Total (QDGCs) 2017/03/24 1803 740 40,865

BODATSA: Species level (QDGCs) 2017/04/05 1803 685 40,139
PHYTOBAS: Species level (GPS) 2018/09/18 3114 373 29,589
BODATSA & PHYTOBAS: Total (QDGCs) 2018/10/26 1811 765 47,652
BODATSA & PHYTOBAS: Species level (QDGCs) 2018/10/26 1803 678 43,889
QDGCs quarter degree grid cells, GPS records with global positioning system localities

PHYTOBAS is a National Vegetation Data Archive
(2003–2009), designed and administered by the late
Dr Bobby Westfall. This database is currently inactive
and the proposal by Specht et al. (2018) that ecological
data should be curated, proposing online open access
of historical data, is supported. Since PHYTOBAS
seldom contain infraspecific taxa, only species level
records were used for further analyses. Introduced
grass species and incomplete taxon records were also
removed. A summary of the extent of the two datasets
following the editing and merging processes is shown
in Table 1. With the exception of references made to
grass pasture species conserved in the SA-NFG, no
other information was sourced from this genebank
database.
The merging of the two datasets resulted in many
duplicated records within a QDGC. After duplicates
were removed, the contributing number of QDGCs for
the two datasets were determined and are shown in
Table 2. BODATSA contributed to 86.7% of the total
dataset and PHYTOBAS 0.7%. Grass species listed by
Fish et al. (2015), but not recorded in either
BODATSA or PHYTOBAS, are documented and thus
the only species not represented in this study. There
were eight indigenous grass species not recorded in
either BODATSA or PHYTOBAS and these are listed
in Table 3.
The collection and/or survey intensity (expressed as
the number of grass species per QDGC collected as
herbarium specimens), henceforth referred to as
collection intensity, was calculated and mapped. This
is also used as a reference map to ascertain the
presence or absence of grass species within QDGCs of
the study area when e.g. distribution maps are
compared.
The taxonomy (subfamily and tribe) and descrip-
tive attributes, specifically endemism, photosynthetic
Table 2 The post-editing contribution of BODATSA and
PHYTOBAS datasets
#QDGC % of Total
BODATSA 1564 86.7
PHYTOBAS 13 0.7
BODATSA & PHYTOBAS merged 226 12.5
Total 1803 100.0
pathway and growth form, were added to each record.
For the purpose of this study, growth forms indicated
as ‘‘climber’’, ‘‘decumbent’’ and ‘‘scrambler’’ were
grouped under the term ‘‘trailing’’. The information
for the various attributes was sourced from, amongst
others, Gibbs Russell et al. (1990), Germishuizen and
Meyer (2003), Van Oudtshoorn (2012), Fish et al.
(2015) and SANBI (2017). The phylogenetic classi-
fication of Soreng et al. (2017) was followed to
compile the evolutionary relationships of subfamilies
and tribes. The bioregions vegetation map of Ruther-
ford et al. (2006) was used as a base layer for
generating maps with ArcView GIS 3.2, ESRI Inc.
2002. This vegetation map shows 35 bioregions where
a bioregion is defined as a composite special terrestrial
unit based on similar biotic (vegetation and floristic)
and physical features (landscapes and rock types) and
processes at the regional scale (Rutherford et al. 2006).
The biomes map of southern Africa (Rutherford et al.
2006) and the Köppen-Geiger climate classification
map of southern Africa (Beck et al. 2018) are Online
Resources 1 and 2 respectively, to be used as reference
maps.
The ‘Red List of South African Plants’ (SANBI
2017), was consulted for data on the conservational
status of indigenous grass species. A list of grasses

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Table 3 List of Poaceae species indigenous to South Africa, Lesotho and Eswatini not recorded in the Botanical Database of
southern Africa (BODATSA; maintained by SANBI) and PHYTOBAS (South African National Vegetation Data Archive) datasets
Species Recorded in
a
Catabrosa drakensbergense (Hedberg & I.Hedberg) Soreng & Fish Lesotho
a
Ellisochloa papposa (Nees) P.M.Peterson & N.P.Barker Eastern Cape
Helictotrichon rogerellisii Mashau, Fish & A.E.van Wyk Western Cape
Helictotrichon roggeveldense Mashau, Fish & A.E.van Wyk Northern Cape
Melinis scabrida (K.Schum.) Hack. Limpopo
Pentameris praecox (H.P.Linder) Galley & H.P.Linder KwaZulu-Natal
Poa leptoclada Hochst. ex A.Rich. KwaZulu-Natal, Lesotho
Tribolium pleuropogon (Stapf) Verboom & H.P.Linder Southern Cape
a
Status: Vulnerable

with categories ‘Rare’ (not exposed to any direct or
potential threat), ‘Near Threatened’ (likely to become
at risk of extinction), ‘Vulnerable’ (high risk of
extinction), ‘Endangered’ (very high risk of extinc-
tion) and ‘Critically Endangered’ (extremely high risk
of extinction) will be presented. The conservation
status, however, applies only to species within South
Africa’s borders and is thus not a global assessment.
Results and discussion
Collection intensity
The collection intensity for indigenous grass species is
shown in Fig. 1. Collection intensity records, as
presented here, are not linked to species diversity as
the latter is determined by vegetation surveys of plant

Fig. 1 Collection intensities of Poaceae indigenous to South SANBI) and PHYTOBAS (South African National Vegetation
Africa, Lesotho and Eswatini as recorded in the Botanical Data Archive) datasets mapped on the bioregions of Rutherford
Database of southern Africa (BODATSA; maintained by et al. (2006)

123

communities. For this study, BODATSA included
records documented since 1802 and PHYTOBAS
records from 2003 to 2009.
It is evident that the very high collection intensities
([ 151 spp./QDGC) were made in the Central
Bushveld bioregion (Rutherford et al. (2006) as shown
in Fig. 2). Moreover, collection efforts were high in
the 2528CA QDGC (red square with [ 201 spp./
QDGC), indicating a possible biased collection
approach close to Pretoria, Gauteng. Collection
intensities ranging from 51 to 150 spp./QDGC are
recorded in the Central Bushveld-, Lowveld-, Indian
Ocean Coastal Belt- and the Fynbos bioregions. The
lower intensities (\ 100 spp./QDGC) are relatively
well distributed over the study area, with no record-
ings, however, in some of the central parts of the arid
region. When the SANBI’s grass herbarium collec-
tions are compared to the legume collections (Tryts-
man et al. 2011), it is evident that the former are larger
(± 40 139 records) than the latter (± 27 618 records)
despite the larger number of indigenous legume
(± 1455 intraspecific taxa of which ± 12% are trees)
versus grass species (± 685 species).
Phylogenetic classification
Worldwide the Poaceae is here treated as containing
12 subfamilies, 52 tribes, 768 genera and 11,506
species (Soreng et al. 2017). The statistics of the
grasses as a whole is compared with the indigenous
grass diversity of SA in Table 4. Grasses, indigenous
to SA, consist of eight subfamilies, 25 tribes, 151
genera and 685 species. The basal lineage subfamilies
Anomochlooideae, Pharoideae and Puelioideae are
not represented in SA. The subfamilies of the BOP
clade (Bambusoideae, Oryzoideae and Pooideae) are
all present in SA, but not Micrairoideae of the
PACMAD clade. The PACMAD clade consist of
subfamilies Panicoideae, Aristidoideae, Chlori-
doideae, Micrairoideae, Arundinoideae and Dantho-
nioideae. Differences in the formal classification of
Poaceae as followed by Gibbs Russell et al. (1990) and
Fish et al. (2015), compared to that of Soreng et al.
(2017), is outlined in Table 5.
Bambusoideae has the poorest representation of
species in SA (Table 4). Given that Bambusoideae is
the only grass lineage to have diversified in forests
(Sungkaew et al. 2009), this is expected with forests
being the smallest biome in the study area (Online
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Genet Resour Crop Evol
Resource 1). Pooideae contains the largest number of
world species (3968 spp.) but Panicoideae the largest
number of species indigenous to SA (256 spp.).
Furthermore, only 20% of the world genera (151
genera) and 6% of world species (685 spp.) are found
in SA. Even though this is a low number of species,
many have proven economic forage value worldwide,
e.g. Anthephora pubescens, Chloris gayana, Digitaria
eriantha and Eragrostis curvula, highlighting the
possible wealth of genetic resources contained in
especially the tribe Paniceae.
As mentioned previously, the largest number of
grass species is contained in Panicoideae resulting
mainly from the high number of species in Paniceae
(150 spp.). Pentameris P.Beauv. is the largest genus
with 75 species followed by Eragrostis Wolf with 65
species (Table 6). Pentameris is found in more
temperate regions, i.e. mainly in the Cape Floristic
Region (Barker 1993), whereas Eragrostis is found in
tropical and subtropical regions (Truter et al. 2015)
(also refer to Online Resource 2 for tropical, subtrop-
ical and temperate regions of SA). The importance of
the Panicoideae is highlighted by the fact that both Zea
mays L. and Sorghum Moench are recognized species
within the Andropogoneae.
Table 6 denotes the genera currently conserved in
the SA-NFG. This genebank conserves at present 73
genera and 162 indigenous grass species, i.e. 48% and
24% of the total number of taxa respectively. The low
number of grass species presently conserved is a
particular concern if the high risk of continued loss of
genetic material as described by Pengelly and Maass
(2018) is taken into account. However, the renewed
interest from South African policy makers in restoring
this genebank as a centre of excellence is encouraging
and strengthen the aim to improve the collection and
conservation efforts for potentially important indige-
nous grass species.
Subfamily distribution
The distribution patterns, i.e. the presence or absence
of subfamilies in QDGCs of the study area, for the
eight subfamilies represented in SA are shown in
Fig. 3. Aristidoideae, Panicoideae and Chloridoideae
covers most of the bioregions with the latter having
more coverage in especially the central regions of
South Africa. The presence of Panicoideae is infre-
quent in Namaqualand, Bushmanland and Karoo
Genet Resour Crop Evol

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b Fig. 2 Bioregions of South Africa, Lesotho and Swaziland Growth form

(Rutherford et al. 2006) to be used as reference map
The majority of grass species in SA are described as
tufted (401 spp.) followed by rhizomatous/tufted (105
regions (refer to Fig. 2) and Aristidoideae has a low spp.) and rhizomatous (59 spp.) (Fig. 4). Linder et al.
presence in the central parts of the arid regions. (2018) states that the tufted growth form, stemming
According to Fish et al. (2015), Panicoideae occurs in from tillering, probably evolved within Poaceae. This
more mesic habitats whereas Chloridoideae occurs in growth form was found, however, to be less tolerant to
drier saline habitats. heavy grazing compared to rhizomatous or stolonif-
erous grasses by Reynolds (1995). The tufted growth
Figure 3 further shows that Oryzoideae and Pooi-
form is nonetheless an attribute of the majority of grass
deae have similar distribution patterns, covering the
pasture species, particularly those mentioned earlier as
Cape Floristic Region, Succulent Karoo and a large
key pasture species. Rhizomes (underground stems)
part of the Grassland biome (Online Resource 1).
are also a valuable survival trait, in not only protecting
Bambusoideae occurs only in the Drakensberg Grass-
the plant from trampling (Rechenthin 1956) or being
land bioregion whereas Arundinoideae occurs mostly
correlated with drought resistance (Zhou et al. 2014)
in Central Bushveld, Lowveld and Indian Ocean
but also having a regenerative ability (Zwerts et al.
Coastal Belt bioregions (refer to Fig. 2). The high
2015). Da Silva et al. (2015) argues that complemen-
presence of Arundinoideae in the winter rainfall zone
tary growth habits of C4 tropical grasses, i.e. rhizoma-
described by Gibbs Russell (1986) has been invali-
tous grasses that occupy a lower horizontal stratum
dated since the addition of the Oryzoideae and
and tufted grasses a higher vertical stratum, result in
Danthonioideae resulted in the removal of Ehrharta
coexistence and allows for greater grass diversity.
Thunb. and Pentameris from Arundinoideae. Lastly,
Little information on the hemicryptophyte growth
Danthonioideae is found in the Cape Floristic Region,
form (buds present at or near the soil surface) exists for
Succulent Karroo biome (Online Resource 1) and the
these indigenous species, although it is accepted that it
Drakensberg Grassland bioregion.

Table 4 Statistics of tribes, genera and species of Poaceae indigenous to South Africa, Lesotho and Eswatini (SA) compared to the
world Sourced from Fish et al. (2015) and Soreng et al. (2017)
Subfamily Tribe World Tribe SA Genera World Genera SA Species World Species SA

Anomochlooideae 2 – 2 – 4 –
Pharoideae 1 – 3 – 12 –
Puelioideae 2 – 2 – 11 –

BOP clade
Oryzoideae 4 2 19 4 115 28
Bambusoideae 3 2 125 2 1670 2
Pooideae 15 6 202 18 3968 58

PACMAD clade
Aristidoideae 1 1 3 3 367 42
Panicoideae 13 6 247 65 3241 256
Arundinoideae 2 2 14 4 40 6
Micrairoideae 3 – 8 – 184 –
Danthonioideae 1 1 19 10 292 114
Chloridoideae 5 5 124 45 1602 179
Total 52 25 768 151 11,506 685

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Table 5 Comparison of the classification of Poaceae indigenous to South Africa, Lesotho and Eswatini followed by Gibbs Russell
et al. (1990) and Fish et al. (2015) compared to that of Soreng et al. (2017)
a b a
Gibbs-Russell et al. (1990) Fish et al. (2015) Soreng et al. (2017)
Subfamily Tribe Subfamily Tribe Subfamily Tribe

Pooideae Triticeae Bambusoideae Bambuseae Oryzoideae Ehrharteae

Brachypodieae Olyreae Oryzeae
Bromeae Ehrhartoideae Ehrharteae Bambusoideae Arundinarieae
Aveneae Oryzeae Bambuseae
Meliceae Pooideae Brachypodieae Pooideae Meliceae
Poeae Bromeae Stipeae
Bambusoideae Oryzeae Meliceae Brachypodieae
Olyreae Poeae Bromeae
Centotheceae Stipeae Triticeae
Ehrharteae Triticeae Poeae
Bambuseae Aristidoideae Aristideae Aristidoideae Aristideae
Arundinoideae Stipeae Arundinoideae Arundineae Panicoideae Tristachyideae
Arundineae Danthonioideae Danthonieae Centotheceae
Danthonieae Panicoideae Andropogoneae Paniceae
Aristideae Arundinelleae Paspaleae
Chloridoideae Pappophoreae Paniceae Arundinelleae
Chlorideae Paspaleae Andropogoneae
Panicoideae Paniceae Tristachyideae Arundinoideae Molinieae
Arundinelleae Centotheceae Arundineae
Andropogoneae Chloridoideae Centropodieae Danthonioideae Danthonieae
Maydeae Cynodonteae Chloridoideae Centropodieae
Eragrostideae Triraphideae
Triraphideae Eragrostideae
Zoysieae Zoysieae
Cynodonteae
a
Tribes in phylogenetical order
b
Tribes in alphabetical order

is common in grasses (Linder et al. 2018). This trait
can contribute to survival during unfavourable condi-
tions such as seasonal drought, cold, fire or heavy
grazing when plants can regrow from the base after
conditions improves (Linder et al. 2018).
In general, most grass growth forms have a
relatively wide distribution throughout the study area.
The exceptions are the cushion growth form (low
growing, mat forming at high altitudes) of the genus
Pentameris (eight of the 75 spp.), the trailing form of
especially eight Panicum L. spp. and the three trailing
Oplismenus P.Beauv. spp. Pentameris is found mainly
in the Fynbos biome, Panicum spp. in the Nama-
Karoo, Grassland, Savanna and Indian Ocean Coastal
Belt biomes and Oplismenus spp. in the Lowveld and
Indian Ocean Coastal Belt bioregions (refer to Fig. 2
and Online Resource 1). A review on the concept of
grazing lawns, i.e. a short-stature grassland commu-
nity type, persisting and spreading under heavy
grazing (Hempson et al. 2014) affirms that certain
ecotypes of Themeda triandra Forssk. and Digitaria
eriantha forms small cushion-like plants and dense
stoloniferous or rhizomatous clonal mats under fre-
quent grazing. This finding strengthens the call of Van
Wyk (1995) to formally label infraspecific genetic

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 Genet Resour Crop Evol

Table 6 Subfamilies, tribes and genera (number of species in phylogenetic classification of Soreng et al. (2017) for subfam-
brackets sourced from Fish et al. (2015)) of Poaceae indige- ilies and tribes. Taxa in bold indicate the largest (most
nous to South Africa, Lesotho and Eswatini following the speciose) group/s within the classification
Subfamily Tribe Genera

BOP clade
Oryzoideae Ehrharteae Ehrhartaab (23)
Oryzeae Leersiab (2) Oryzab (2) Prosphytochloaa (1)
Bambusoideae Arundinarieae Thamnocalamusa (1)
Bambuseae Oxytenanthera (1)
i
Pooideae Meliceae Melicaa (2) Streblochaete (1)
Stipeae Stipaa (2)
Brachypodieae Brachypodiuma (2)
Bromeae Bromusa (4)
Triticeae Hordeuma (1) Secalea (1)
Poeae Agrostisa (9) Anthoxanthuma (4) Calamagrostis (1) Catabrosa (1) Festucaa (8) Holcusa (1)
Helictotrichona (14) Koeleria (1) Poab (3) Polypogona (1) Puccinelliaa (2)
PACMAD clade
Aristidoideae Aristideae Aristidaab (23) Sartidia (2) Stipagrostisab (17)
Panicoideae Tristachyideae Danthoniopsisb (5) Loudetiab (5) Trichopteryx (1) Tristachyab (3)
Centotheceae Megastachya (1)
Paniceaecef Acrocerasb (1) Alloteropsisb (2) Anthephoraab (3) Brachiariab (16) Cenchrusb (1) Digitariaab
(25) Echinochloab (9) Entolasia (1) Eriochloab (4) Leucophrys (1) Megaloprotachneb (1)
Melinisab (8) Odontelytrum (1) Oplismenus (3) Panicumab(34) Paspalidiumb (2)
Pennisetum (7) Pseudechinolaena (1) Sacciolepis (6) Setariaab (12) Stenotaphrumb (2)
Stereochlaena (1) Tarigidiaa (1) Tricholaenaab (2) Urochloab (6)
Paspaleae Paspalumb (3)
Arundinelleae Arundinella (1)
Andropogoneaeeg Andropogonb (14) Arthraxon (1) Bothriochloab (3) Chrysopogonb (1) Cleistachneb (1)
Coelorachis (1) Cymbopogonab (6) Dichanthiumb (1) Diheteropogonb (2) Elionurusb (1)
Elymandra (1) Eriochrysis (2) Eulalia (2) Hackelochloa (1) Hemarthriab (1) Heteropogonb
(2) Hyparrheniab (20) Hypertheliab (1) Imperatab (1) Ischaemumb (2) Miscanthusb (2)
Monocymbiumb (1) Oxyrhachis (1) Phacelurus (1) Rhytachne (2) Rottboellia (1)
Schizachyriumb (6) Sehima (2) Sorghastrumab (2) Sorghumb (2) Themedab (1)
Trachypogonb (1) Urelytrum (1)
Arundinoideae Molinieae Elytrophorus (1) Phragmites (2) Styppeiochloa (1)
Arundineae Dregeochloaa (2)
Danthonioideae Danthonieae Capeochloaa (3) Chaetobromusab (1) Geochloaa (3) Merxmuelleraa (4) Pentamerisa (75)
Pentaschistisa (3) Pseudopentamerisa (3) Schismusab (4) Tenaxiaa (5) Triboliuma (13)
Chloridoideae Centropodieae Centropodiaab (1) Ellisochloa (1)
Triraphideae Triraphisa (4)
Eragrostideaedh Catalepis (1) Cladoraphisa (2) Diandrochloa (2) Enneapogonab (6) Eragrostisab (65)
Fingerhuthiaab (2) Pogonarthriab (1) Schmidtiab (2) Stiburus (2) Tetrachneb (1)
Zoysieae Spartina (1) Sporobolusab (34)

123
Genet Resour Crop Evol

Table 6 continued
Subfamily Tribe Genera

Cynodonteaec Acrachneb (1) Bewsiab (1) Brachychloa (2) Chlorisb (6) Coelachyrum (1) Ctenium (1)
Cynodonab (6) Dactylocteniumb (4) Dinebra (1) Eleusineb (2) Enteropogonb (2) Eustachysb
(1) Harpochloa (1) Leptocarydion (1) Leptochloab (5) Lepturusb (1) Lintoniab (1)
Lophacmea (1) Microchloab (2) Mosdeniaa (1) Odyssea (1) Oropetiumb (1) Perotis (1)
Polevansia (1) Rendlia (1) Schoenefeldiab (1) Tetrapogonb (1) Tragusab (3) Trichoneurab
(2) Tripogon (1)
a
Include endemic species
b
Genera conserved in the South African National Forage Genebank
Tribes with majority of species with cPioneer, dSubclimax, eClimax, fDecreaser, gIncreaser I, hIncreaser II
i
Temperate grass lineage

variants within especially species of potential eco-
nomic significance.
Photosynthetic pathway and grazing status
The photosynthetic pathway distinguished in the
subfamilies and tribes of Poaceae is shown in Table 7.
The early-branching subfamilies (Oryzoideae, Bam-
busoideae and Pooideae) are shown to have a C3
photosynthetic pathway, whereas the later-branched
Chloridoideae has a C4 photosynthetic pathway.
Interestingly, Bouchenak-Khelladi et al. (2010) found
molecular evidence that C4 photosynthesis (of at least
the subfamily Chloridoideae) may well have origi-
nated in Africa. The majority of species of Aristi-
doideae and Panicoideae are C4 whereas all species of
Arundinoideae and Danthonioideae are C3. Tribes
with well-known pasture species i.e. Andropogoneae,
Eragrostideae and Cynodonteae contains only C4
species whereas Paniceae have a mixture of C3 (i.e.
genus Panicum) and C4 species. The importance of C4
grasses in pasture production could lie in their ability
to transfer larger proportions of plant nitrogen to roots
in infertile environments (Long 1999), as well as
larger leaf area production in fertile and disturbed
environments. This supports Linder et al. (2018)
finding that grasses has the ability to colonize, persist
and transform environments, properties that are the
key to success. Furthermore, evidence presented by
Linder et al. (2018) indicated that C4 plants have a
higher carbon fixing efficiency over a range of habitats
when soil resources are limited compared to C3 types.
However, the higher forage value in terms of crude
protein content and digestibility in C3 compared to C4
grasses (Gibson 2009) could in effect be an important
grass collection objective, where the focus is on
pasture development for temperate regions, especially
for dairy farming.
The distribution of C3 and C4 grass species is shown
in Fig. 5. Species with the C3 pathway are widespread
in the Fynbos, Succulent Karoo biomes and the
Drakensberg Grassland, Sub-Escarpment Grassland
bioregions (refer to Fig. 2 and Online Resource 1).
The central arid region has low occurrences of C3
grass species whereas the Kalahari Duneveld biore-
gion has none. Species with a C4 photosynthetic
pathway is found in all the bioregions of SA. Vogel
et al. (1978) investigated the geographical distribution
of C3 and C4 grasses in southern Africa and concluded
that low temperatures during seasonal growth favour
the C3 grasses in the regions mentioned above. The
exclusive presence of C4 in the tropical region (Online
Resource 2) as described by Vogel et al. (1978) is,
however, not evident in Fig. 5. C3 species such as
Agrostis L., Festuca L., Helictotrichon Besser (Poeae)
and Pentameris (Danthonieae) were recorded in the
tropical regions of the study area (Online Resource 2).
The only known grass species in SA with both a C3 and
C4 subspecies, Alloteropsis semialata (R.Br.) Hitchc.
need special mentioning (Ellis 1974; Gibbs Russell
1983). Alloteropsis semialata (R.Br.) Hitchc. subsp.
eckloniana (Nees) Gibbs Russ. has a C3 photosyn-
thetic pathway and A. semialata (R.Br.) Hitchc. subsp.
semialata (R.Br.) Hitchc. a C4, suggesting an evolu-
tionary reversion from C4 to C3 (Ibrahim et al. 2009).
Within Panicum, a genus containing important forage
crops, Ellis (1988) distinguished 11 Panicum spp.
having a C3 and 23 with C4 photosynthetic pathway.

123
 Genet Resour Crop Evol

Fig. 3 The distribution patterns for Poaceae subfamilies indigenous to South Africa, Lesotho and Eswatini in phylogenetic order
according to Soreng et al. (2017), mapped on the bioregions of Rutherford et al. (2006)

123
Genet Resour Crop Evol

450

400 401

350
Number of grass species

300

250

200

150

105
100
59
50
18
1… 13 12 8 8 6 6 6 4 3 3 3 3 3 2 1 1
0

Hydrophyte/trailing
Stoloniferous

Stoloniferous/tufted

Cushion/tufted
Tufted

Cushion

Hydrophyte/stoloniferous/tufted

Tufted/trailing

Geophytic
Trailing

Rhizomatous/stoloniferous

Hydrophyte/rhizomatous/stoloniferous

Hydrophyte/tufted

Rhizomatous/stoloniferous/tufted

Hydrophyte/rhizomatous

Hydrophyte/rhizomatous/stoloniferous/tufted

Rhizomatous/woody
Hydrophyte

Hydrophyte/rhizomatous/tufted
Rhizomatous
Rhizomatous/tufted

Fig. 4 Growth forms of Poaceae indigenous to South Africa, Lesotho and Eswatini. Sourced mainly from Gibbs Russell et al. (1990)

Pau et al. (2012) pointed out that the evolutionary Andropogoneae are well presented in all biomes but
history of Poaceae is important for understanding the narrowly presented in the drier areas namely the
C3 and C4 functional diversity of grasses, as this will Nama-Karoo and Succulent Karoo biomes (Online
affect their responses to global change. Resource 1). The ecological status as defined by
A comparison of the recorded successional status of Forani et al. (1978) and Van Oudtshoorn (2012)
tribes (Van Oudtshoorn 2012) shows that Paniceae and indicate that the high number of Decreaser species
Cynodonteae contain the largest number of species present in Paniceae further underlines this tribe’s
with pioneer status, whereas Paniceae and Andro- forage value, i.e. 47% of indigenous species with
pogoneae contain the largest number with climax known preferential grazing status are grouped here
status (Table 7). Pioneer species are usually annuals, (Table 7). Decreasers are defined as grasses abundant
growing in disturbed habitats or unfavourable condi- in good veld and will decrease when over- or
tions whereas climax species are perennials, growing undergrazed, whereas Increasers will increase under
only when normal, optimal growth conditions prevail any type of mismanagement. Andropogoneae and
(Van Oudtshoorn 2012). Panicoideae distinctly con- Eragrostideae contain respectively the highest number
tains most climax species compared to other subfam- of Increaser I and Increaser II species.
ilies and thus should be the focus for further assessing
grass genetic resources with pasture potential, espe-
cially within the tribes Paniceae and Andropogoneae.
Grass species belonging to Paniceae and

123
 Genet Resour Crop Evol

Table 7 The number of species within Poaceae subfamilies from Gibbs Russell et al. (1990) and successional and
and tribes indigenous to South Africa, Lesotho and Eswatini ecological grazing status (Van Oudtshoorn 2012)
using a C3 and/or C4 photosynthetic pathway sourced mainly
Subfamily Tribe C3 C4 Pioneer Subclimax Climax Decreaser Increaser Increaser Increaser
I II III

BOP clade
Oryzoideae Ehrharteae 23 2 1 2
Oryzeae 5
Bambusoideae Arundinarieae 1
Bambuseae 1
Pooideae Meliceae 3 1 1
Stipeae 2
Brachypodieae 2
Bromeae 4 1
Triticeae 2
Poeae 45 1 1 2 2 2

PACMAD clade
Aristidoideae Aristideae 2 40 4 3 5 2 9 3
Panicoideae 14 2 1
Tristachyideae
Centotheceae 1
Paniceae 23 127 13 3 25 21 7 16
Paspaleae 3 1
Arundinelleae 1 1 1
Andropogoneae 87 5 27 7 20 5 2
Arundinoideae Molinieae 4 1
Arundineae 2
Danthonioideae Danthonieae 114 1 3 3 1 2 2
Chloridoideae Centropodieae 2 1
Triraphideae 4 1 1
Eragrostideae 84 7 11 5 2 1 25 2
Zoysieae 35 2 3 2 1 1 6 1
Cynodonteae 54 10 3 6 5 2 14 1

Endemism and conservation concern
Figure 6 shows the collection intensity for the 257
grass species endemic to SA. The highest number of
endemic species per QDGC is recorded in the Cape
Floristic Region and in the Drakensberg Alpine Centre
(Van Wyk and Smith 2001; Rutherford et al. 2006). In
terms of the presence of endemic species in the study
area, the Upper and Lower Karoo, Bushmanland and
Lowveld are the main bioregions (refer to Fig. 2)
containing the smallest number of endemic species.
Danthonioideae (found mainly in the Cape Floristic
Region and Succulent Karoo biome (see Online
Resource 1) contributes to nearly half of the total
number of endemic grass species in SA. Pentameris
accounts for 28% of endemic species followed by
Ehrharta (9%), Eragrostis (7%) and Tribolium (5%).
Pentameris is mainly found in the Fynbos biome and
Drakensberg Grassland bioregion, Ehrharta in the
Fynbos and Succulent Karoo biome, Eragrostis in
most parts of the study area and Tribolium in the
Fynbos and Succulent Karoo biome (Fish et al. 2015).

123
Genet Resour Crop Evol
Fig. 5 The distribution of C3 and C4 photosynthetic pathways
in Poaceae indigenous to South Africa, Lesotho and Eswatini.
Sourced mainly from Gibbs Russell et al. (1990) and mapped on
the bioregions of Rutherford et al. (2006)
Pentameris and Tribolium are considered having a
lower grazing value than Ehrharta (Van Oudtshoorn
2012).
The conservational assessment for grass species
published in the Red List of South African Plants
(SANBI 2017) is listed in Table 8 together with
regions or centres of endemism (sensu Van Wyk
and Smith 2001). The genus Pentameris holds a
true conservational concern with 24 species listed,
present in all categories. There are two Panicum
spp. and one Secale L. sp., both important genera
in pasture production also listed, respectively as
rare and critically endangered. Panicum sancta-
luciense and Panicum dewinteri, both perennials,
are found respectively in the tropical region
(Online Resource 2) and the Mopane bioregion
of the study area. Secale strictum (J.Presl) J.Presl
subsp. africanum (Stapf) K.Hammer, is a peren-
nial wild rye found in Renosterveld of the
western mountain Karoo (Fish et al. 2015) and
within the Hantam-Roggeveld Centre (Van Wyk
and Smith 2001). Helictotrichon quinquesetum
(Steud.) Schweick., only known from the slopes
of Table Mountain, Cape Town (Fish et al.
2015), is the only grass species listed as possibly
extinct. It is evident from Table 8 that the
majority of grass species with a conservational
concern are found in the Cape Floristic Region,
followed by the Hatam–Roggeveld Centre.
Habitat loss is identified as the major threat to South
African plants, i.e. infrastructure development, urban
expansion, cultivation of crops, commercial afforesta-
tion and mining (SANBI 2017). In terms of the
conservation of genetic resources for possible future
use, it is suggested that representative seed samples
are collected from these species and stored in the SA-
NFG. The screening and characterisation of these
genetic resources will allow for re-introduction when
certain populations go locally extinct. In addition,
these efforts will allow these genetic resources to be
included into breeding programs that can produce
more adapted species or for rehabilitation of degraded
rangelands.
Conclusion
The important role that members of the Poaceae plays
in sustainable pasture production systems compels the
SA-NFG to conserve SA’s indigenous grass genetic
resources, not only those used in current pasture
systems, but also genetic material that could be
beneficial in future breeding programs and/or adapted
to specific agro-ecological conditions. A modern
classification framework was therefore used to docu-
ment indigenous grass species, recorded in various
southern African taxonomic reviews and botanical
databases. Linking various attributes with phyloge-
netic classification and vegetation types assisted in
distinguishing taxa with pasture potential and to
prioritise future collection and conservation efforts.
123

Fig. 6 The collection intensity for the endemic grass species present in South Africa, Lesotho and Eswatini. Sourced from Fish et al.
(2015) and mapped on the bioregions of Rutherford et al. (2006)
Results showed that the inclusion of PHYTOBAS
(South African National Vegetation Data Archive)
added valuable data since 13 additional QDGCs were
added to the dataset. Attention is drawn to the value of
historical vegetation data and the proposed call for
online open access is supported. The collection
intensity map shows that the central Bushveld has
high collections, but that in large parts of the central
arid region no collections were done. There are eight
subfamilies, 25 tribes, 151 genera and 685 species
present in SA. Subfamilies Anomochlooideae, Phar-
oideae, Puelioideae (basal lineage) and Micrairoideae
(PACMAD clade) are not represented in SA. Only 6%
of world grass species (i.e. 685 spp.) are found in SA,
with Panicoideae having the largest number of species
(256 spp.). Aristidoideae, Panicoideae and Chlori-
doideae are represented in most of the bioregions
whereas Bambusoideae is found only in one bioregion,
namely the Drakensberg Grassland. The infrequent
presence of Panicoideae in the western and central arid
regions should be taken into account when focusing on
the collection of potentially drought tolerant grass
species. Instead, the presence of Chloridoideae in
123
Genet Resour Crop Evol
these arid regions, containing important pasture
species such as Chloris, Cynodon and Eragrostis,
should be considered. The majority of grass species in
SA are tufted, an attribute of the majority of key grass
pasture species. Tribes with well-known pasture
species contains only C4 species (Andropogoneae,
Eragrostideae and Cynodonteae) whereas Paniceae
have a mixture of C3 and C4 species (i.e. genus
Panicum). Species with a C4 photosynthetic pathway
is found in all the bioregions of SA whereas C3 species
have low occurrences in the central arid region.
Panicoideae contains more climax species compared
to other subfamilies, especially within the tribes
Paniceae and Andropogoneae. The 257 endemic grass
species found largely in the Cape Floristic Region and
in the Drakensberg Alpine Centre need an in depth
assessment to determine the role the SA-NFG plays in
their conservation as possible future pasture genetic
resources. This is an urgent outcome for the 24 species
of Pentameris that holds a true conservational con-
cern. The collection of viable seed of two Panicum
spp. and one Secale sp., listed respectively as rare and
critically endangered and the possible extinct Secale
Genet Resour Crop Evol

Table 8 Grass species and infraspecific taxa on the ‘Red List
of South African Plants’ in order of the least to highest risk of
extinction (SANBI 2017). The superscript following some
names indicates to which specific local region or centre of
Rare
endemism (sensu Van Wyk and Smith 2001) the particular
taxon is endemic, or if more widespread, confined to in the
study area
Vulnerable (continued)

Capeochloa setaceaCFR Elytrophorus globularis

Dregeochloa calviniensis Helictotrichon barbatumSKR
Panicum sancta-lucienseMC Helictotrichon namaquenseKBC
CFR
Pentameris caulescens Helictotrichon rogerellisiiCFR
CFR
Pentameris clavata Oryza longistaminata
Pentameris glacialisCFR Pentameris calcicola var. hirsutaCFR
CFR
Pentameris hirtiglumis Pentameris longiglumis subsp. longiglumisCFR
CFR
Pentameris holciformis Pentameris longipesAC
Pentameris horridaCFR Pentameris trifidaCFR
CFR
Pentameris longiglumis subsp. gymnocolea Sartidia jucundaSBC
CFR
Pentameris swartbergensis Tribolium ciliareCFR
Pentameris unifloraCFR
Near threatened Endangered

Eulalia aurea Helictotrichon roggeveldenseHRC

PC
Oxyrhachis gracillima * Pentameris bachmanniiCFR
SBC
Panicum dewinteri Pentameris dentataHRC
CFR
Pentameris aspera Pentameris eckloniiCFR
Pentameris calcicola var. calcicolaCFR Pentameris pholiuroidesCFR
Pentameris limaKBC Pentameris scandensCFR
Stipagrostis geminifoliaGC

Vulnerable Critically endangered

Capeochloa cincta subsp. sericeaAC Pentameris barbata subsp. orientalisCFR

DAC
Catabrosa drakensbergense Pentameris elegansCFR
Ehrharta setacea subsp. unifloraCFR Pentameris ellisiiCFR
CFR
Ellisochloa papposa Secale strictum subsp. africanumHRC
Tribolium pleuropogonCFR
Helictotrichon quinquesetum (possibly extinct)CFR
Rare, not exposed to any direct or potential threat; Near Threatened, likely to become at risk of extinction; Vulnerable, high risk of
extinction; Endangered, very high risk of extinction; Critically Endangered, extremely high risk of extinction; AC, Albany Centre;
CFR, Cape Floristic Region; DAC, Drakensberg Alpine Centre; GC, Gariep Centre; HRC, Hantam–Roggeveld Centre; KBC,
Kamiesberg Centre; MC, Maputaland Centre; PC, Pondoland Centre; SBC, Soutpansberg Centre; SKR, Succulent Karoo Region
*In the study area confined to the centre, but not endemic as it also occurs elsewhere in Africa/Madagascar

strictum subsp. africanum is a critical undertaking for
the SA-NFG.
The SA-NFG conserve at present only a quarter of
indigenous grass species with the current collection
already facing serious risks. This will prompt decision
makers to reinvest in the SA-NFG, focusing on
conserving genetic resources for improved animal
production, mitigate the possible effect of climate
change on the valuable grass genetic resources, and
consequently pasture production. It is proposed that a
strategy be developed for the SA-NFG to collect and
conserve seed of Paniceae since this tribe contains
valuable pasture grass species, thus focusing on
speciose subtropical genera such as Anthephora,
Brachiaria, Digitaria, Panicum and Setaria.

123

Furthermore, temperate genera containing important
pasture species i.e. Bromus, Hordeum and Festuca as
well as endemic species with grazing value, such as
Ehrharta, should also be the focus of seed collection
efforts by the SA-NFG. The need for more emphasis
on the description and formal recognition of
infraspecific taxa, including ecotypes, are also high-
lighted. It is further proposed that stakeholders in
biodiversity conservation strategize plant collections
excursions to those areas, previously not sampled.
The current systems of germplasm conservation at
the SA-NFG is under revision. A long term storage
facility is being prepared for purely conservation
purposes, while active collections of important pasture
species as well as indigenous species with agronomic
potential will be made available for research purposes.
The reason for the long term storage of all grass
genetic resources was brought about by the changes in
bioclimatic conditions predicted for southern Africa.
The conservation of these resources at the SA-NFG
could, under future bioclimatic conditions, result in
potential rehabilitation of degraded areas, or reintro-
ductions with better adapted ecotypes of the same
species after local extinction of naturally occurring
populations. The indigenous grass database developed
for this study will be used to establish biogeographical
patterns of the grass flora as well as for assessing their
pasture potential. These results will be combined with
the published results for indigenous legumes and used
to develop a collective strategy in terms of prioritizing
species, identifying key regions and planning charac-
terization and evaluation studies.
Acknowledgements We thank the South African National
Biodiversity Institute (SANBI) for making available the
distribution and descriptive data contained in the BODATSA
database, the late Dr Bobby Westfall for administrating the data
contained in the PHYTOBAS National Vegetation Data Archive
and Elsa van Niekerk (ARC-PPR) for the graphics.
Funding The Agricultural Research Council of South Africa
funded this study.
Data availability The datasets generated during and/or
analysed during the current study are available from the
corresponding author on reasonable request.
123
Genet Resour Crop Evol
Compliance with ethical standards
Conflict of interest The authors declare that they have no
conflict of interest.
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