Check List
Check List 13(1): 2030, Erratum, 24 January 2017
ISSN 1809-127X © 2017 Check List and Authors
LISTS OF SPECIES
Erratum
Woody species of the Miombo woodlands and geoxylic
grasslands of the Cusseque area, south-central Angola
Rasmus Revermann, Francisco Maiato Gonçalves, Amândio Luis Gomes & Manfred Finckh
In Table 1, on page 7, please read “Oleaceae” in place of “Olacaceae”. The authors regret this error.
The original, unaltered publication begins on the following page.
i
the journal of
biodiversity data
Check List
the journal of
biodiversity data
Check List 13(1): 2030, 6 January 2017 doi: https://doi.org/10.15560/13.1.2030
ISSN 1809-127X © 2017 Check List and Authors
LISTS OF SPECIES
Woody species of the Miombo woodlands and geoxylic
grasslands of the Cusseque area, south-central Angola
Rasmus Revermann1, 4, Francisco Maiato Gonçalves1, 2, Amândio Luis Gomes1, 3 & Manfred Finckh1
1
University of Hamburg, Biocentre Klein Flottbek, Department of Biodiversity, Ecology and Evolution of Plants, Ohnhorststr. 18, 22609
Hamburg, Germany
2
Herbarium of Lubango, ISCED-Huíla, Department of Natural Sciences, Rua Sarmento Rodrigues, 230 Lubango, Angola
3
University Agostinho Neto, Faculty of Sciences, Department of Biology, Av. 4 de Fevereiro, Luanda, Angola
4
Corresponding author. E-mail: rasmus.revermann@uni-hamburg.de
types and over 100 subordinate types dealt with in the text
(Barbosa 1970, 1971). However, the descriptions of these
vegetation types were of general character and limited to
the dominant species. Detailed descriptions of the species
composition and plant diversity of the Angolan vegetation
are lacking for most parts of the country. For the province
of Bié, Monteiro (1970) provided an excellent overview
on the woody vegetation including the first provincial
map of the woodlands. Based on 144 relevés, Monteiro
delineated three associations of woody plants and four
sub-associations.
The civil war that followed Angola’s independence in 1975
made any scientific work in Angola extremely difficult, and
thus, most scientific literature available today dates back to
the pre-independence era. Since the end of the armed conflict in 2002, scientific work is slowly increasing. However,
botanical work in the country is still hampered by the lack
of field guides and the fact that the principal work on the
flora of Angola, the Conspectus Florae Angolensis, remains
unfinished and important families such as the Rubiaceae
are not treated. Similarly, a countrywide checklist of the
flora of Angola was lacking until the recent publication of
“Plants of Angola – Plantas de Angola” by Figueiredo &
Smith (2008). Nevertheless, inventories of vascular plants
at the local and regional scale are still lacking for most
parts of the country. Such inventories are indispensable
for any kind of natural resource management planning,
conservation measures or ecological studies (Figueiredo
et al. 2009).
The interdisciplinary research project “The Future
Okavango” (TFO) aims to provide a scientific basis for
strategic resource planning for the Okavango Basin. The
headwaters of the Okavango River, where 95% of the
runoff are generated, are located on the Angolan Central
Plateau (Steudel et al. 2013). Rapid transformations of
the social-ecological systems are currently taking place
there (Pröpper et al. 2015). However, little knowledge
abstract: The species composition of the vegetation in
most regions of Angola has been poorly studied and most
studies date back to the pre-independence era. In this
study, we provide a detailed account of the woody flora
of the Miombo woodlands and geoxylic grasslands of
the Cusseque study site of “The Future Okavango” (TFO)
project, situated on the Angolan Central Plateau. The
checklist is based on a vegetation survey using vegetation
plots of 1,000 m2 and also includes records from botanical
collections made elsewhere at the study site. In total, we
documented 154 woody species belonging to 99 genera of
37 plant families in 100 km2. The study represents the first
comprehensive account of the woody vegetation of the
area including all habitats and growth forms.
Key words: Angola; Bié; geoxylic suffrutex; Miombo; The Future
Okavango project; vegetation survey
INtrODuCtION
Rural communities in Angola hold an enormous knowledge
of the local flora and especially have great understanding
of the potential usages of plants (Figueiredo & Smith
2012; Kissanga 2016). In contrast, scientific exploration and documentation of the vegetation of Angola is
still limited. Early botanists such as Friedrich Welwitsch
visited the country in the middle of the 19th century (Welwitsch 1869). The most influential botanist working in
Angola in the first half of the 20th century was John Gossweiler, who worked in all Angolan provinces and collected
over 14,000 specimens. His collection is considered an
especially important source of information for rare and
endemic species (Figueiredo & Smith 2008). Furthermore, Gossweiler produced the first phytogeographic map
of Angola containing 19 principal vegetation types (Gossweiler & Mendonça 1939). Based on this map and his
own observations, Luís A. Grandvaux Barbosa published
a new phytogeographic map in 1971 containing 32 main
1
Revermann et al. | Woody species of the Cusseque area, south-central Angola
season lasting from November to April. The mean annual
precipitation is 987 mm and the mean annual temperature
is 20.4°C (Weber 2013). The study area harbours a high
pedodiversity. The elevated areas are characterized by
deep and developed slightly loamy Arenosols. The slopes
of the smaller valleys of the tributaries and at the western
side of the Cusseque River show shallow Plinthisols on
granitic bedrock. The soils along the eastern part of the
Cusseque River are characterized by very deep and leached
Arenosols. The centre of the valleys support Histosols with
peat layers exceeding 1 m in depth while at the edges of the
wetlands Gleysols are the common soil type (Gröngröft
et al. 2013a).
The main vegetation types covering south central
Angola are semi-deciduous Miombo woodlands and forests
(Figures 2a and b). These woodlands are interspersed with
open vegetation types locally termed anharas de ongote.
The salient feature of the open vegetation types are
dwarf shrubs with a huge underground woody biomass.
This distinct life form was described by White (1976)
as “geoxylic suffrutex”. In the Cusseque area, geoxylic
suffrutices occur on two different soil types: on deep,
leached sandy soils and on shallow, compact, ferralitic
soils. Accordingly, we will differentiate herein between
“geoxylic grasslands on sandy soils” (Figure 2c) and
“geoxylic grasslands on ferralitic soils” (Figure 2d). The
occurrences of the different vegetation types are governed
by topography: woodlands and forests are confined to
the elevated areas and upper slopes. The mid- and lower
and data on the vegetation and the botanical diversity was
available (Revermann 2016). In this study, we present
results of the vegetation survey carried out at the research
site “Cusseque” in the province of Bié located at the upper
reaches of the Okavango River.
matErIaLS aND mEtHODS
Study site
The Okavango River originates on the Angolan Central
Plateau and terminates in a large inland delta in the
Kalahari Desert in Botswana. Within the TFO project,
detailed studies were carried out at four research sites
representing the different parts of the river basin. The
work presented in this paper was carried out at the
study site Cusseque with an area of 100 km2 (13.6985° S,
017.0382° E). The site is located on the Angolan Central
Plateau in the province of Bié (Figure 1; Wehberg &
Weinzierl 2013). The landscape can be described as a
rolling plain intersected by the Cusseque River and its
many tributaries, which are orientated perpendicular to
each other. Three major landscape units can be identified:
the elevated areas, the sloping areas leading down to the
valley bottoms and the valley floors (Gröngröft et al.
2013b). The mean elevation is 1,575 m above sea level
while the difference in elevation between the valley bottom
of the main river and the surrounding elevated areas is
about 100 m (Gröngröft et al. 2013b). The climate of
the Cusseque area is semi-humid with a pronounced wet
Figure 1. Location of the Okavango Basin in southern Africa and the study site “Cusseque” denoted in red. (Projection: WGS 1984; background: RapidEye
high-resolution satellite imagery, recorded 1 May 2013. We acknowledge the DLR for the provision of the data from the RapidEye Science Archive.)
Check List | www.biotaxa.org/cl
2
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
Figure 2. Landscapes of the Cusseque study area: a) Miombo woodland in the middle of the rainy season, b) Miombo woodland at the end of the rainy
season, c) geoxylic grasslands dominated by Cryptosepalum maraviense at the beginning of the rainy season in October, d) geoxylic grassland on sandy
soils at the beginning of the dry season in May; in the background wetland on the valley bottom and Miombo woodlands.
slopes feature geoxylic grasslands. The woodlands and
geoxylic grasslands are separated by ecotones extending
up to several hundred meters where elements of both
vegetation types co-occur. The valley bottoms support
wetlands dominated by Cyperaceae (Revermann et al.
2013; Schneibel et al. 2013).
to the unique character of the geoxylic grasslands, these
were subject to an additional field study. Therein, data were
collected using 10 m × 10 m plots with two 3.3 m × 3.3 m
subplots situated in diagonally opposite corners (adapted
from Dengler 2009). In every plot all vascular plants
found were recorded and their projected cover estimated
visually. Unknown plants were photographed and voucher
specimens were collected according to botanical methods
outlined by Victor et al. (2004). Voucher specimens were
deposited in the herbarium of the ISCED Huíla (LUBA)
and in the Herbarium Hamburgense (HBG). In addition to
the species recorded on the vegetation plots, species found
elsewhere while working at the study area were added to
the checklist. A high number of vegetation plots and several months of field work carried out by four observers in all
seasons ensured a comprehensive coverage of the woody
species present at the study area.
Data collection
Plot based vegetation surveys were carried out during the
growing season in the years 2011 to 2014 and all information is stored in the Vegetation Database of the Okavango
Basin (GIVD ID: AF-00-009, Revermann et al. 2016). In
order to evenly map all existing vegetation units, sampling
followed a random, stratified design. Based on an image
segmentation algorithm using all bands of a Landsat 7
scene, seven major vegetation units were identified. In
these vegetation units random points were created using
GIS and transferred to a hand-held GPS for localization in
the field. Furthermore, additional vegetation plots were
examined in different successional stages of Miombo forest to analyse successional pathways of the regeneration of
natural vegetation after disturbance by shifting cultivation
(Gonçalves et al., accepted). We used a nested plot design
with a 10 m × 10 m plot located in the centre of a 20 m ×
50 m plot. In total, we sampled 148 vegetation plots. Due
Check List | www.biotaxa.org/cl
taxonomy and plant identification
We followed the taxonomy of the checklist “Plants of Angola – Plantas de Angola” by Figueiredo & Smith (2008). We
are aware of recent changes in the taxonomy, but decided
to conform to the national checklist. For identification,
we consulted the Conspectus Florae Angolensis (Exell &
Mendonça 1937, 1951, 1954, 1955; Exell & Fernandes
3
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
1962, 1966; Exell et al. 1970) when possible, and the flora
of neighbouring countries, especially the Flora Zambeziaca
(Exell & Wild 1960) and the field guide to the Trees and
shrubs of Namibia (Mannheimer & Curtis 2009). Additionally, we consulted herbarium collections at the ISCED
Huíla (LUBA) and the Instituto de Investigação Científica
Tropical (LISC) as well as the on-line database JSTOR Plant
Science (http://plants.jstor.org/). For some specimens, we
consulted experts at Kew Botanical Garden (K).
Permits
Permits for plant collection and transfer of biological
material in Angola for scientific purposes was arranged
based on the framework of Material Transfer Agreements
from Angola, negotiated between the Instituto Superior
de Ciências de Educação da Huíla (ISCED, Huíla), Lubango
and the University of Hamburg (UHH), Germany and
authorized on behalf of the Angolan Government by the
Director for Agriculture, Fisheries and Environment of the
Province of Huíla. All International Conventions to which
Angola is signatory country, such as Convention on International Trade in Endangered Species of Wild Fauna and
Flora (1973), Convention on Biological Diversity (1992),
International Treaty on Plant Genetic Resources for Food
and Agriculture (2004), and all other relevant national and
international instruments concerning biodiversity were
taken into account.
Figure 3. The families of woody plants found at the Cusseque study
site, south-central Angola. Only families with more than two species are
shown. In total 37 families of woody plant species occurred, containing
154 species in 99 genera.
and forests showed the highest woody species richness with
110 species belonging to 32 families. A surprisingly high
number of 33 woody species from 14 families were found in
the geoxylic grasslands. These open vegetation types appear
to be merely grasslands in the late growing season (Figures
2c and d) but in fact harbour a remarkable diversity of woody
species (Table 1). In the wetlands few woody species occurred
and we recorded only two Ficus species sporadically occurring along the margins of the wetlands.
One species, Combretum schumannii Engl., was recorded
but is not listed in the current checklist of Angola
(Figueiredo & Smith 2008).
Data analysis
DISCuSSION
We visualized the number of species per family of all
species belonging to one family using the function ‘pie’ in
the statistical software R (R Development Core Team
2016). Data on the frequency of occurrence of a species, the
habitat and the life form were compiled from vegetation
plot data. We assigned frequency according to the following
categories: very rare (1 or 2 observations), rare (3–5
observations), occasional (6–10 observations), frequent
(11–30 observations), common (>30 observations). We
assigned every species to one or more of the following life
form categories based on field observations and literature:
tree, shrub, liana, dwarf shrub and geoxyle. For geoxyles
we followed the definition proposed by White (1976).
White defined a geoxyle as a dwarf shrub that has closely
related species growing as trees and that exhibits massive
woody underground parts.
The 154 species recorded within the 100 km2 of the Cusseque
study site almost equalled the 166 woody species found by
Monteiro (1970) in the entire province of Bié, an area of
70,314 km2. This does not reflect the quality of the study of
Monteiro but rather illustrates how poorly the region has
been surveyed so far. In fact, the study carried out by Monteiro (1970) is of high quality and stands out as the only
study of its time from Angola basing its analysis on quantitative, plot based data. However, the study was restricted to
woodlands and only larger shrubs and trees were included.
In contrast, we included all vegetation types ranging from
woodlands to geoxylic grasslands and wetlands. Due to
the high sampling intensity and the coverage of all vegetation types our species list can therefore be considered a
comprehensive checklist of the woody plant species of the
Cusseque area. However, it must be noted that this list does
not contain any specimen that could not be identified to
at least genus level; some specimens were lacking fruits or
flowers, preventing further identification. Therefore, the
actual number of woody species may be slightly higher.
Despite the relatively recent publication of the checklist
of vascular plants of Angola, subsequent field surveys in
various parts of the country have resulted in additions to
the checklist (Huntley & Coelho 2011). We provided the
first record in Angola of the herbaceous Asteraceae Schistostephium crataegifolium (DC.) Fenzl ex Harv, during the
rESuLtS
We documented 154 woody species belonging to 99 genera
and 37 families (Table 1). The majority of species belonged to
the family Fabaceae (33), followed by Rubiaceae (22), Euphorbiaceae (11), Proteaceae (10) and Combretaceae (9) (Figure
3). Most of the dominant species belonged to the Fabaceae
and occurred with high frequencies. In contrast, the Rubiaceae, second in species richness, contained species occurring
with low frequencies and were less abundant. The woodlands
Check List | www.biotaxa.org/cl
4
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
Table 1. List of species arranged by family. Voucher specimens have been deposited in the herbaria of Lubango (LUBA) and Hamburg (HBG). In most
cases doublets are stored in both herbaria. The herbarium name in parentheses is the location where the specimen used for identification is deposited.
Frequency was assigned according to the following categories: very rare (1 or 2 observations), rare (3–5 observations), occasional (6–10 observations),
frequent (11–30 observations), common (>30 observations). Life forms of the species were assigned to one or more of the categories: tree, shrub, liana,
dwarf shrub and geoxyle.
Species name
Frequency
Life form
Habitat
Collection number(s)
Anacardiaceae
Ozoroa cf. xylophylla (Engl. &Gilg) R.Fern. & A.Fern.
very rare
shrub
geoxylic grassland (sandy soils)
133057B (HBG)
Ozoroa stenophylla Engl. & Gilg.
frequent
shrub
woodland / grassland (ferralitic
soils)
140123 (LUBA)
Rhus arenaria Torre, A.R.
frequent
dwarf shrub /
geoxyle
geoxylic grassland (ferralitic
soils)
140101 (LUBA)
Rhus exelliana Meikle
frequent
dwarf shrub
Rhus gracilipes Exell
frequent
dwarf shrub
woodland / forest
139227; 132483 (HBG)
135250; 134275 (HBG)
Rhus kirkii Oliv.
frequent
dwarf shrub
woodland / forest
139253 (LUBA)
Anisophylleaceae
Anisophyllea boehmii Engl.
frequent
tree
woodland / forest
134316; 139018; 135297 (HBG)
Anisophyllea quangensis Engl. ex Henriq.
rare
dwarf shrub
geoxylic grassland (sandy soils)
133044; 134116 (HBG); 140109 (LUBA)
Annoncaceae
Annona stenophylla ssp. nana Engl. & Diels
rare
dwarf shrub
woodland / forest
133058; 134218 (HBG); 140065 (LUBA)
Uvaria angolensis Welw. ex Oliv. (Figure 4b)
frequent
shrub
woodland / forest
135323; 134240 (HBG)
Xylopia odoratissima Welw. ex Oiv.
frequent
shrub
woodland / forest
133057A; 134263 (HBG)
Xylopia tomentosa Exell
common
shrub
woodland / forest
135279; 132956; 132986 (HBG); 139177
(LUBA)
common
dwarf shrub /
geoxyle
geoxylic grassland (sandy soils)
140121 (LUBA)
Apocynaceae
Chamaeclitandra henriquesiana (Hallier f.) Pichon
Diplorhynchus condylocarpon (Müll. Arg.) Pichon
frequent
shrub / tree
woodland / forest
135300 (HBG)
Landolphia camptoloba (K.Schum.) Pichon
frequent
liana
woodland / forest
132537 (HBG)
Landolphia gossweileri (Stapf ) Pichon
rare
dwarf shrub
geoxylic grassland (sandy soils)
133048 (HBG)
Strophanthus welwitschii (Baill.) K.Schum.
frequent
liana
woodland / forest
135336; 135378; 134091 (HBG)
Asparagaceae
Asparagus sp. 135286
frequent
shrub
woodland / forest
135286 (HBG)
Asparagus cf. africanus Lam.
very rare
shrub
woodland / forest
134115 (HBG)
occasional
shrub
woodland / forest
132695 (HBG)
Parinari capensis Harv.
frequent
dwarf shrub /
geoxyle
geoxylic grassland (sandy soils)
132664; 132898; 140068 (HBG)
Parinari curatellifolia Planch. ex Benth.
common
tree
woodland / forest
132444 (HBG)
Asteraceae
Helichrysum krausii Sch. Bip
Crysobalanaceae
Combretaceae
Combretum acutifolium Exell
very rare
liana / shrub
woodland / forest
135306 (HBG)
Combretum collinum Fresen.
common
tree
woodland / forest
139176 (HBG)
Combretum elaeagnoides Klotzsch
very rare
tree
woodland / forest
132538 (HBG)
Combretum engleri Schinz
frequent
shrub
woodland / forest
133216 (HBG)
Combretum platypetalum ssp. platypetalum Welw.
ex M.A.Lawson (Figure 4f )
occasional
dwarf shrub
geoxylic grassland (sandy soils)
132639; 134114; 140113 (HBG)
Combretum schumannii Engl.
rare
shrub
woodland / forest
139048 (LUBA)
Combretum zeyheri Sond.
frequent
shrub / tree
woodland / forest
135280; 132510 (HBG)
Pteleopsis anisoptera (Welw.) Engl. & Diels
frequent
shrub / tree
woodland / forest
135365; 134110 (HBG); 139066 (LUBA)
Terminalia brachystemma Welw. ex Hiern
frequent
tree
woodland / forest / grassland
(sandy and ferralitic soils)
132997; 134088; 134131 (HBG)
frequent
dwarf shrub /
geoxyle
geoxylic grassland (sandy soils)
140165 (LUBA)
Dichapetalaceae
Dichapetalum cymosum (Hook.) Engl.
Dipterocarpaceae
Monotes africanus A.DC.
common
tree
woodland / forest
132917; 134160; 134228 (HBG)
Monotes angolensis de Wild.
very rare
tree
woodland / forest
132443 (HBG)
Monotes caloneurus Gilg.
rare
tree
woodland / forest
134820 (HBG)
Monotes dasyanthus Gilg
common
tree
woodland / forest
132907; 132961 (HBG); 139228 (LUBA)
Ebenaceae
Diospyros batocana Hiern
occasional
shrub / tree
woodland / forest
139247 (LUBA)
Diospyros chamaethamnus Dinter ex Mildbr.
frequent
dwarf shrub /
geoxyle
geoxylic grassland (sandy soils)
140179 (LUBA)
Continued
Check List | www.biotaxa.org/cl
5
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
Table 1. Continued.
Species name
Frequency
Life form
Habitat
Collection number(s)
Ebenaceae, continued
Diospyros pseudomespilus ssp. brevicalyx Mildbr.
Diospyros virgata (Gürke) Brenan
Euclea crispa ssp. crispa (Thunb.) Gürke
frequent
occasional
frequent
shrub
shrub
dwarf shrub
woodland / forest
woodland / forest
woodland / forest / geoxylic
grassland
135379 (HBG)
132941 (HBG)
135413 (HBG)
very rare
shrub / tree
woodland / forest
139235 (LUBA)
occasional
frequent
rare
rare
common
common
occasional
frequent
common
shrub / tree
shrub / tree
shrub / tree
dwarf shrub
dwarf shrub
dwarf shrub
tree
dwarf shrub
dwarf shrub /
geoxyle
tree
tree
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
geoxylic grassland (ferralitic
soils)
woodland / forest
woodland / forest
139095 (LUBA)
134099; 134135 (HBG); 139068 (LUBA)
139113 (LUBA)
139256 (LUBA)
139238 (LUBA)
139237 (LUBA)
132555; 134232 (HBG); 139038 (LUBA)
132990; 134185 (HBG)
132490; 134199 (HBG)
132691; 132912; 132998 (HBG)
geoxylic grassland (sandy soils)
140167 (LUBA)
frequent
occasional
frequent
common
frequent
dwarf shrub /
geoxyle
tree
shrub / tree
shrub / tree
shrub
shrub / tree
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
134156; 135318 (HBG); 139223 (LUBA)
139065; 135342; 132967 (HBG)
135360; 139242; 133018 (HBG)
135311 (HBG)
132963; 139128 (LUBA)
common
occasional
common
common
common
rare
rare
rare
common
tree
tree
tree
tree
shrub
dwarf shrub
dwarf shrub
dwarf shrub
tree
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
135298; 139016 (LUBA)
132957; 139255 (LUBA)
132676 (HBG)
132900; 135335 (HBG); 139233 (LUBA)
139121 (LUBA)
139257 (LUBA)
139196 (LUBA)
135304 (HBG); 139023 (LUBA)
common
dwarf shrub
132754; 132825 (HBG)
common
dwarf shrub
Dalbergia nitidula Welw. ex Baker
Dialium englerianum Henriq.
Dolichos sp. 140088
rare
frequent
frequent
Entada arenaria Schinz
Eriosema sp. 133109
very rare
rare
shrub / tree
shrub / tree
dwarf shrub /
geoxyle
dwarf shrub
dwarf shrub
Eriosema sp. 132895
rare
dwarf shrub
Erythrina abyssinica Lam. ex DC.
Erythrophleum africanum (Welw. ex Benth.) Harms
Guibourtia coleosperma (Benth.) J.Léonard
Humularia welwitschii (Taub.) P.A.Duvign.
Indigofera baumiana Harms
Indigofera congesta Welw. ex Baker
Kotschya strobilantha (Welw. ex Baker) Dewit & P. A.
Duvign. var. strobilantha
Mucuna sp. 140052
very rare
common
occasional
common
frequent
occasional
rare
140052 (LUBA)
Pericopsis angolensis (Baker) Meeuwen
Pterocarpus angolensis DC
frequent
occasional
tree
tree
tree
dwarf shrub
shrub
dwarf shrub
dwarf shrub /
geoxyle
dwarf shrub /
geoxyle
shrub / tree
tree
geoxylic grassland (ferralitic
soils)
geoxylic grassland (ferralitic
soils)
woodland / forest
woodland / forest
geoxylic grassland (ferralitic
soils)
geoxylic grassland (sandy soils)
geoxylic grassland (ferralitic
soils)
geoxylic grassland (ferralitic
soils)
giant termite mounds
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
geoxylic grassland (ferralitic
soils)
geoxylic grassland (sandy soils)
woodland / forest
woodland / forest
139181 (LUBA)
-
Ericaceae
Erica benguellensis (Welw. ex Engl.) E.G.H. Oliv.
Euphorbiaceae
Bridelia sp. 139095
Hymenocardia acida Tul.
Maprounea africana Müll. Arg.
Phyllanthus angolensis Müll. Arg.
Phyllanthus sp. 139238
Phyllanthus welwitschianus Müll. Arg.
Pseudolachnostylis maprouneifolia Pax
Sclerocroton oblongifolius (Müll. Arg.) Kruijt & Roebers
Uapaca sp. 134199
Uapaca kirkiana Müll. Arg.
Uapaca nitida var. nitida Müll. Arg.
Fabaceae
Abrus melanospermus ssp. suffruticosus (Boutique)
D.K.Harder
Albizia antunesiana Harms
Albizia gummifera (J.F.Gmel) C.A.Sm.
Baphia bequaertii De Wild.
Bauhinia petersiana Bolle
Bobgunnia madagascariensis (Desv.) J.H.Kirkbr. &
Wiersema
Brachystegia bakeriana Hutch. & Burtt Davy
Brachystegia longifolia Benth.
Brachystegia spiciformis Benth.
Burkea africana Hook.
Copaifera baumiana Harms
Crotalaria amoena Welw. ex Baker
Crotalaria cistoides Welw. ex Baker
Crotalaria florida Welw. ex Baker
Cryptosepalum exfoliatum ssp. pseudotaxus (Baker f.)
P.A.Duvign. & Brenan (Figure 4i)
Cryptosepalum exfoliatum ssp. suffruticans
(P.A.Duvign.) P.A.Duvign. & Bre (Figure 4h)
Cryptosepalum maraviense Oliv. (Figure 4g)
common
common
occasional
frequent
135308B; 135620 (HBG)
139236 (LUBA)
133147; 139034 (LUBA)
140088 (LUBA)
134147 (HBG)
133109 (HBG)
132753; 132895 (HBG)
135333 (HBG)
139054 (LUBA)
139146 (LUBA)
132530 (HBG)
139237 (LUBA)
139141 (LUBA)
Continued
Check List | www.biotaxa.org/cl
6
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
Table 1. Continued.
Species name
Frequency
Life form
Habitat
Collection number(s)
Hypericaceae
Psorospermum febrifugum Spach.
rare
shrub / tree
woodland / forest
139036 (LUBA)
Psorospermum tenuifolium Hook.f.
rare
shrub / tree
woodland / forest
132958 (HBG)
common
shrub
woodland / forest
132968; 133005; 133149 (HBG)
Ixonanthaceae
Phyllocosmus lemaireanus (De Wild. & T.Durand)
T.Durand & H.Durand
Lamicaceae
Alvesia rosmarinifolia Welw.
occasional
shrub
woodland / forest
134776; 132533 (HBG)
Tinnea sp. 133121
frequent
dwarf shrub
geoxylic grassland (ferralitic
soils)
133121 (HBG)
Vitex doniana Sweet
occasional
shrub
woodland / woodland ecotone
132915 (HBG)
Vitex madiensis Oliv.
frequent
shrub
woodland / woodland ecotone
132996 (HBG); 139069 (LUBA)
Melastomataceae
Memecylon flavovirens Baker
frequent
shrub / tree
woodland / forest
132519; 133161 (HBG); 139240 (LUBA)
Warneckea sapinii (De Wild.) Jacq.-Fél. (Figure 4d)
occasional
tree
woodland / forest
135309 (HBG); 139140 (LUBA)
occasional
shrub
woodland / forest
132546; 133000; 133096 (HBG)
Meliaceae
Ekebergia benguelensis Welw. ex C.DC.
Moraceae
Ficus pygmaea Welw. ex Hiern
rare
dwarf shrub
wetland margin
141510 (HBG)
Ficus sp. 141539
rare
dwarf shrub
Wetland margin
141539 (HBG)
rare
dwarfshrub /
geoxyle
geoxylic grassland (sandy soils) 140118 (LUBA)
common
shrub
woodland / forest
Myricaceae
Morella cf. serrata (Lam.) Killick
Myrsinaceae
Myrsine africana L.
134107; 134278 (HBG); 139024 (LUBA)
Myrtaceae
tree
woodland / forest
135813 (HBG)
Syzygium guineense ssp. macrocarpum (Engl.) F.White common
Syzygium guineense ssp. barotsense F.White
occasional
shrub / tree
woodland ecotone
135800; 135796 (HBG)
Syzygium guineense ssp. huillense (Hiern) F.White
frequent
dwarf shrub
geoxylic grassland (sandy soils)
133072; 135614; 135882 (HBG)
Ochna afzelii ssp. mechowiana R.Br. ex Oliv.
rare
dwarf shrub /
geoxyle
woodland / grassland
133128 (HBG)
Ochna arenaria De Wild. & T.Durand (Figure 4e)
frequent
dwarf shrub
woodland, geoxylic grassland
(sandy and ferralitic soils)
132947; 133024 (HBG); 140016 (LUBA)
Ochnaceae
Ochna manikensis De Wild.
frequent
dwarf shrub
geoxylic grassland (sandy soils)
132654; 132803 (HBG)
Ochna pulchra Hook.
common
shrub / tree
woodland / forest
135381; 139064 (LUBA)
Ochna pygmaea Hiern
common
dwarf shrub /
geoxyle
woodland / forest, grassland
(sandy soils)
139239; 140154 (LUBA)
Olacaceae
Jasminum pauciflorum Benth.
rare
liana / shrub
woodland / forest
139238 (LUBA)
Schrebera trichoclada Welw.
rare
shrub / tree
woodland / forest
139189 (LUBA)
rare
dwarf shrub
woodland / forest
139033 (LUBA)
common
shrub
woodland / forest
135299 (HBG); 139242 (LUBA)
rare
shrub / tree
woodland / forest
139208 (LUBA)
occasional
tree
woodland / forest
133017 (HBG)
frequent
shrub
woodland / forest
135322; 133032 (HBG); 139164 (LUBA)
Orobanchaceae
Sopubia karaguensis Oliv.
Passifloraceae
Paropsia brazzaeana Baill.
Picodendraceae
Oldfieldia dactylophylla (Welw. ex Oliv.) J.Léonard
Polygalaceae
Securidaca longepedunculata Fresen
Polygonaceae
Oxygonum fruticosum Dammer ex Milne-Redh.
Proteaceae
Faurea intermedia Engl. & Gilg
occasional
shrub / tree
woodland / forest
132720; 139072 (LUBA)
Faurea rochetiana (A.Rich.) Chiov. ex Pic.Serm.
frequent
tree
woodland / forest / ecotone
135307 (HBG)
Faurea saligna Harv.
occasional
tree
geoxylic grassland (ferralitic
soils)
132549; 132980; 134205 (HBG)
Protea baumii Engl. &Gilg.
occasional
dwarf shrub
woodland / forest
132501; 133019; 134225 (HBG)
Protea gaguedi J.F.Gmel.
frequent
tree
woodland / forest
132918 (LUBA)
Protea angolensis var. divaricarta (Engl. & Gilg.) Beard
rare
dwarf shrub
geoxylic grassland (ferralitic
soils)
134200 (HBG)
Protea micans ssp. trichophylla Welw.
occasional
dwarf shrub
geoxylic grassland (sandy soils)
132607 (HBG); 140096 (LUBA)
Proteaceae, continued
Continued
Check List | www.biotaxa.org/cl
7
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
Table 1. Continued.
Species name
Life form
Habitat
Collection number(s)
Protea petiolaris ssp. petiolaris (Hier) Baker & C.H.Wright frequent
Frequency
tree
woodland / forest
132982 (HBG)
Protea cf. welwitschii Engl.
rare
dwarf shrub
geoxylic grassland
(ferraliticsoils)
132480 (HBG)
Protea sp. 133045
rare
dwarf shrub
geoxylic grassland (sandy soils)
133045 (HBG)
rare
shrub
woodland / forest
133093; 135285 (HBG)
rare
dwarf shrub
geoxylic grassland (on sandy
soils) & woodland ecotone
136003 (HBG)
Rhamnaceae
Ziziphus mucronata Willd.
Rubiaceae
Ancylanthos rubiginosus Desf.
Fadogia cf. chrysantha K.Schum.
very rare
shrub
woodland ecotone
134257 (HBG)
Fadogia cf. triphylla var. triphylla Baker
very rare
shrub
woodland / forest
132987 (HBG); 133081 (HBG)
Fadogia fuchsioides Welw. ex Oliv. (Figure 4c)
occasional
shrub
woodland / forest
132524 (HBG)
Fadogia cf. homblei De Wild.
rare
dwarf shrub /
geoxyle
geoxylic grassland
140114 (LUBA)
Fadogia cf. monticola Robyns
rare
dwarf shrub /
geoxyle
geoxylic grassland
140146 (LUBA)
Fadogia sp. 134097
occasional
shrub
geoxylic grassland (ferralitic
soils), woodland ecotone
132453; 134097; 134167 (HBG)
135338 (HBG)
Gardenia brachythamnus (K.Schum.) Launert
very rare
dwarf shrub
woodland ecotone
Keetia cf. gracilis (Hiern) Bridson
very rare
shrub
woodland / forest
132442; 133148 (HBG)
Keetia venosa (Oliv.) Bridson
rare
shrub
woodland / forest
132534A (HBG)
Leptactina benguellensis (Welw. ex Benth. & Hook.f.)
R.D.Good
rare
dwarf shrub
woodland / forest
135313; 135353; 133153 (HBG)
Leptactina prostrata K.Schum
very rare
dwarf shrub
geoxylic grassland (ferralitic
soils)
134181 (HBG)
Pachystigma pygmaeum (Schltr.) Robyns
frequent
dwarf shrub /
geoxyle
geoxylic grassland (sandy soils)
140138 (LUBA)
Pygmaeothamnus cf. chamaedendrum (Kuntze) Robyns very rare
dwarf shrub
geoxylic grassland (sandy soils)
132723 (HBG)
Pygmaeothamnus sp. 132552
dwarf shrub
woodland / forest
132552 (HBG)
very rare
Pygmaeothamnus zeyheri (Sond.) Robyns
rare
dwarf shrub
geoxylic grassland (sandy soils)
132798; 133033; 134089 (HBG)
Rytigynia orbicularis (K.Schum.) Robyns
frequent
shrub
woodland / forest
132925; 134127 (HBG); 139056 (LUBA)
Tapiphyllum cf. psammophilum (S.Moore) Robyns
very rare
shrub
woodland / forest
134279 (HBG)
Tricalysia angolensis A.Rich. ex DC.
very rare
shrub
woodland / forest
132500; 133012 (HBG)
Tricalysia sp. 134221
rare
shrub
woodland / forest
134221 (HBG)
Tricalysia coriacea ssp. nyassae (Benth.) Hiern
occasional
shrub
woodland / forest
133008; 134095; 134170 (HBG)
Tricalysia sp. 135367
very rare
shrub
woodland / forest
135367 (HBG)
rare
dwarf shrub
woodland / forest
139228 (LUBA)
Santalaceae
Thesium sp. 139228
Sapotaceae
Chrysophyllum bangweolense R.E.Fr.
rare
tree
woodland / forest
135359 (HBG)
Englerophytum magalismontanum (Sond.) T.D.Penn.
common
shrub
woodland / forest
135320 (HBG); 133151 (HBG); 139109
(LUBA)
rare
shrub
woodland / forest
135308A (HBG)
Smilacaceae
Smilax anceps Willd.
Strychnaceae
Strychnos cocculoides Baker
frequent
shrub / tree
woodland / forest
139070 (LUBA)
Strychnos pungens Soler.
common
shrub / tree
woodland / forest
139254 (LUBA)
Strychnos spinosa Lam.
occasional
tree
woodland / forest
135301 (HBG)
field work for this study (Gonçalves et al. 2016). In the
case of Combretum schumannii Engl. literature indicated
that the range of the species might extend to Angola
(Exell & Wild 1960). However, neither this taxon nor
its synonyms were included in the Angolan checklist
(Figueiredo & Smith 2008).
Most of the species occurred either in the geoxylic grasslands or in the woodlands and forests. However, many of the
geoxylic suffrutices have closely related tree species growing
nearby in the woodlands (Figures 4h and 4i). In Africa, the
centre of diversity of geoxylic suffrutices is in the Zambezian phytoregion. In regions with similar environmental
Check List | www.biotaxa.org/cl
conditions, such as the Sudanian phytoregion, there is only
a very limited number of geoxylic species (White 1976).
We found two types of geoxylic grasslands, each harbouring a very distinct species pool with only a small overlap.
There is much debate on the environmental factors driving
the emergence of this distinct life form (Davy 1922; White
1976; Maurin et al. 2014; Finckh et al. 2016). However, the
different species composition of the two types of geoxylic
grasslands found in Cusseque can be clearly attributed to the
contrasting edaphic conditions. The two dominant species
in the “geoxylic grasslands on ferralitic soils” Cryptosepalum
maraviense (Figure 4g) and C. exfoliatum ssp. suffruticans
8
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
Figure 4 Typical plants of the Cusseque area: a) Copaifera baumiana, b) Uvaria angolensis, c) Fadogia fuchsioides, d) Warneckea sapinii, e) Ochna arenaria,
f) Combretum platypetalum ssp. platypetalum, g) Cryptosepalum maraviense, h) Cryptoseplaum exfoliatum ssp. suffruticans, i) Cryptosepalum exfoliatum
ssp. pseudotaxus.
(Figure 4i) belong to the Fabaceae. In contrast, the “geoxylic grasslands on sandy soils” were dominated by various
species of the genus Ochna of the Ochnaceae and Parinari
capensis of the Chrysobalanaceae. The “geoxylic grasslands
on ferralitic soils” have their core distribution on the Angolan Central Plateau and make up 8.5% of the land surface
within the Cubango Basin (Revermann et al. in revision).
The “geoxylic grasslands on sandy soils” have a very limited
distribution within the study site and cover 0.7% of the area
of the Cubango Basin. However, they are more extensive
further east in the Cuito River Basin and in eastern Moxico
Province, where they occur on large sandy, alluvial plains of
the Zambezi Graben, e.g., in Cameia National Park.
LItEraturE CItED
Barbosa, L.A.G. 1970. Carta fitogeográfica de Angola. Luanda: Instituto de Investigação Científica de Angola. 323 pp.
Barbosa, L.A.G. 1971. Phytogeographical map of Angola. Mitteilungen der Botanischen Staatssammlung München 10: 114–115.
http://biodiversitylibrary.org/page/15185175
Davy, B.J. 1922. The suffrutescent habit as an adaptation to environment. Journal of Ecology 10: 211–219. http://www.jstor.org/
stable/2255742
Dengler, J. 2009. A flexible multi-scale approach for standardised
recording of plant species richness patterns. Ecological Indicators
9: 1169–1178. doi: 10.1016/j.ecolind.2009.02.002
Exell, A.W. & Fernandes, A. 1962. Conspectus Florae Angolensis.
Lisbon: Junta de Investigações do Ultramar. Vol. 3, Fasc. 1: 1–187.
Exell, A.W. & A. Fernandes. 1966. Conspectus Florae Angolensis.
Lisbon: Junta de Investigações do Ultramar. Vol. 3, Fasc. 2:
189–408.
Exell, A.W., A. Fernandes & E.J. Mendes. 1970. Conspectus Florae
Angolensis. Lisbon: Junta de Investigações do Ultramar and
Instituto de Investigação Científica de Angola. Vol. 4: 401 pp.
Exell, A.W. & F.A. Mendonça. 1937b. Conspectus Florae Angolensis.
Lisbon: Junta de Investigações Colonias. Vol. 1, Fasc. 1: 1–176.
Exell, A.W. & F.A. Mendonça. 1951. Conspectus Florae Angolensis.
Lisbon: Junta de Investigações do Ultramar. Vol. 1, Fasc. 2:
177–422.
Exell, A.W. & F.A. Mendonça. 1954. Conspectus Florae Angolensis.
Lisbon: Junta de Investigações do Ultramar. Vol. 2, Fasc. 1: 1–152.
Exell, A.W. & F.A. Mendonça. 1955. Conspectus Florae Angolensis.
aCKNOWLEDGEmENtS
Research was funded by the German Federal Ministry of
Education and Research (BMBF) in the context of The Future
Okavango (TFO) project, grant number 01LL0912A. We are
grateful for the support of the staff at Kew Royal Botanical
gardens who aided in the identification, in particular David
J. Goyder and Iain Darbyshire. Furthermore, we thank the
people of the villages Kaololo, Sovi, Cusseque and Calomba
and especially the traditional authorities (Sobas) for their
support of our study.
Check List | www.biotaxa.org/cl
9
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
ommendations for action. Research project final synthesis report
2010–2015. Hamburg: Universtiy of Hamburg, Biocentre Klein
Flottbek. 190 pp.
R Development Core Team. 2016. R: A language and environment
for statistical computing. https://www.R-project.org/
Revermann, R. 2016. Analysis of vegetation and plant diversity patterns in the Okavango basin at different spatial scales — integration
of field based methods, remote sensing information and ecological
modelling [Dissertation]. Hamburg: University of Hamburg. 295
pp. http://ediss.sub.uni-hamburg.de/volltexte/2016/8156/pdf/
Dissertation.pdf
Revermann, R., A.L. Gomes, F.M. Gonçalves, F. Lages & M.
Finckh. 2013. Cusseque — vegetation. Biodiversity and Ecology
5: 59–63. doi: 10.7809/b-e.00247
Revermann, R., A.L. Gomes, F.M. Gonçalves, J. Wallenfang, T.
Hoche, et al. 2016. Vegetation database of the Okavango Basin.
Phytocoenologia 46: 103–104. doi: 10.1127/phyto/2016/0103
Revermann, R., J. Oldeland, F.M. Gonçalves, J. Luther-Mosebach, A.L. Gomes, et al. [In review]. Dry tropical forests of the
Cubango basin in southern Africa — a first classification and
assessment of their woody species diversity. Phytocoenologia.
Schneibel, A., M. Stellmes, D. Frantz, M. Finckh & R. Revermann. 2013. Cusseque — earth Observation. Biodiversity and
Ecology 5: 55–57. doi: 10.7809/b-e.00246
Steudel, T., H. Göhmann, W.-A- Flügel & J. Helmschrot. 2013.
Assessment of hydrological dynamics in the upper Okavango
River Basins. Biodiversity and Ecology 5: 247–261. doi: 10.7809/
b-e.00279
Victor, J.E., M. Koekemoer, L. Fish, S.J. Smithies & M. Mössmer.
2004. Herbarium essentials: the Southern African Herbarium
user manual. Pretoria: SABONET. 93 pp.
Weber, T. 2013. Cusseque — climate. Biodiversity and Ecology 5:
45–46. doi: 10.7809/b-e.00243
Wehberg, J. & T. Weinzierl. 2013. Okavango Basin — physicogeographical setting. Biodiversity and Ecology 5: 11–13. doi: 10.7809/
b-e.00236
Welwitsch, F. 1869. Sertum Angolense, sive stirpium quarundam
novarum vel minus cognitarum. Transactions of the Linnean
Society 27: 1–94. doi: 10.1111/j.1096-3642.1869.tb00202.x
White, F. 1976. The underground forests of Africa: a preliminary
review. The Gardens’ Bulletin Singapore 11: 57–71.
Lisbon: Junta de Investigações do Ultramar. Vol. 2, Fasc. 2:
153–320.
Exell, A.W. & H. Wild. 1960. Flora Zambesiaca. Glasgow: University
Press Glasgow. 336 pp.
Figueiredo, E. & G.F. Smith. 2008. Plants of Angola — Plantas de
Angola, Strelitzia. Pretoria: South African National Biodiversity
Institute. 279 pp.
Figueiredo, E., G.F. Smith & J. César. 2009. The flora of Angola:
first record of diversity and endemism. Taxon 58: 233–236.
http://www.jstor.org/stable/27756837
Figueiredo, E. & G.F. Smith. 2012. Common names of Angolan
plants. Pretoria: Inhlaba Books. 262 pp.
Finckh, M., R. Revermann & M.P.M. Aidar. 2016. Climate refugees
going underground — a response to Maurin et al. 2014. New Phytologist 209: 904–909. doi: 10.1111/nph.13567
Gonçalves, F.M.P., R. Revermann, A.L. Gomes, M.P.M. Aidar,
M. Finckh & N. Jürgens. [Accepted]. Tree species diversity and
composition of Miombo woodlands in south-central Angola,
a chronosequence of forest recovery after shifting cultivation.
International Journal of Forestry Research.
Gonçalves, F.M., J.J. Tchamba & D.J. Goyder. 2016. Schistostephium
crataegifolium Compositae: Anthemideae, a new generic record.
Bothalia — African Biodiversity and Conservation 46: a209. doi:
10.4102/abc.v46i1.202 9
Gossweiler, J. & F.A. Mendonça. 1939. Carta fitogeográphica de
Angola. Lisbon: República Portuguesa Ministério das Colónias.
242 pp.
Gröngröft, A., J. Luther-Mosebach, L. Landschreiber & A.
Eschenbach. 2013a. Cusseque — soils. Biodiversity and Ecology
5: 51–54. doi: 10.7809/b-e.00245
Gröngröft, A., J. Luther-Mosebach, L. Landschreiber, R. Revermann, M. Finckh & A. Eschenbach. 2013b. Cusseque — landscape.
Biodiversity and Ecology 5: 43–44. doi: 10.7809/b-e.00242
Huntley, B.J. & P. Coelho. 2011. Avaliação Rápida Da Biodiversidade
Da Região Da Lagoa de Carumbo/Rapid Biodiversity Assessment
of the Carumbo Lagoon Area. Luanda: Ministério do ambiente,
República de Angola. 219 pp.
Kissanga Vicente da Silva Firmino, R. 2016. Valorização da flora
de Cusseque e Caiúndo no centro e sul de Angola e avaliação
da biomassa lenhosa utilizada para combustível e construção
[Master’s thesis]. Lisbon: University of Lisbon. 63 pp.
Mannheimer, C. & B.A. Curtis. 2009. Trees and shrubs of Namibia.
Windhoek: Macmillan Education Namibia. 526 pp.
Maurin, O., T.J. Davies, J.E. Burrows, B.H. Daru, K. Yessoufou, et
al. 2014. Savanna fire and the origins of the ‘underground forests’
of Africa. New Phytologist 204: 201–214. doi: 10.1111/nph.12936
Monteiro, R.F.R. 1970. Estudo da flora e da vegetação das florestas
abertas do plantalto do Bié. Luanda: Instituto de Investigação
Científica de Angola. 352 pp.
Pröpper, M., A. Gröngröft, M. Finckh, S. Stirn, V. De Cauwer,
et al. 2015. The Future Okavango — findings, scenarios and rec-
Check List | www.biotaxa.org/cl
authors’ contributions: All authors contributed to the manuscript,
carried out fieldwork and worked on the identification of the
collected specimens. RR wrote the initial draft of the manuscript,
analysed the data and compiled the figures and tables.
received: 19 December 2015
accepted: 7 December 2016
academic editor: Alexander Zizka
10
Volume 13 | Issue 1 | Article 2030
Check List
the journal of
biodiversity data
Check List 13(1): 2030, 6 January 2017 doi: https://doi.org/10.15560/13.1.2030
ISSN 1809-127X © 2017 Check List and Authors
LISTS OF SPECIES
Woody species of the Miombo woodlands and geoxylic
grasslands of the Cusseque area, south-central Angola
Rasmus Revermann1, 4, Francisco Maiato Gonçalves1, 2, Amândio Luis Gomes1, 3 & Manfred Finckh1
1
University of Hamburg, Biocentre Klein Flottbek, Department of Biodiversity, Ecology and Evolution of Plants, Ohnhorststr. 18, 22609
Hamburg, Germany
2
Herbarium of Lubango, ISCED-Huíla, Department of Natural Sciences, Rua Sarmento Rodrigues, 230 Lubango, Angola
3
University Agostinho Neto, Faculty of Sciences, Department of Biology, Av. 4 de Fevereiro, Luanda, Angola
4
Corresponding author. E-mail: rasmus.revermann@uni-hamburg.de
types and over 100 subordinate types dealt with in the text
(Barbosa 1970, 1971). However, the descriptions of these
vegetation types were of general character and limited to
the dominant species. Detailed descriptions of the species
composition and plant diversity of the Angolan vegetation
are lacking for most parts of the country. For the province
of Bié, Monteiro (1970) provided an excellent overview
on the woody vegetation including the first provincial
map of the woodlands. Based on 144 relevés, Monteiro
delineated three associations of woody plants and four
sub-associations.
The civil war that followed Angola’s independence in 1975
made any scientific work in Angola extremely difficult, and
thus, most scientific literature available today dates back to
the pre-independence era. Since the end of the armed conflict in 2002, scientific work is slowly increasing. However,
botanical work in the country is still hampered by the lack
of field guides and the fact that the principal work on the
flora of Angola, the Conspectus Florae Angolensis, remains
unfinished and important families such as the Rubiaceae
are not treated. Similarly, a countrywide checklist of the
flora of Angola was lacking until the recent publication of
“Plants of Angola – Plantas de Angola” by Figueiredo &
Smith (2008). Nevertheless, inventories of vascular plants
at the local and regional scale are still lacking for most
parts of the country. Such inventories are indispensable
for any kind of natural resource management planning,
conservation measures or ecological studies (Figueiredo
et al. 2009).
The interdisciplinary research project “The Future
Okavango” (TFO) aims to provide a scientific basis for
strategic resource planning for the Okavango Basin. The
headwaters of the Okavango River, where 95% of the
runoff are generated, are located on the Angolan Central
Plateau (Steudel et al. 2013). Rapid transformations of
the social-ecological systems are currently taking place
there (Pröpper et al. 2015). However, little knowledge
abstract: The species composition of the vegetation in
most regions of Angola has been poorly studied and most
studies date back to the pre-independence era. In this
study, we provide a detailed account of the woody flora
of the Miombo woodlands and geoxylic grasslands of
the Cusseque study site of “The Future Okavango” (TFO)
project, situated on the Angolan Central Plateau. The
checklist is based on a vegetation survey using vegetation
plots of 1,000 m2 and also includes records from botanical
collections made elsewhere at the study site. In total, we
documented 154 woody species belonging to 99 genera of
37 plant families in 100 km2. The study represents the first
comprehensive account of the woody vegetation of the
area including all habitats and growth forms.
Key words: Angola; Bié; geoxylic suffrutex; Miombo; The Future
Okavango project; vegetation survey
INtrODuCtION
Rural communities in Angola hold an enormous knowledge
of the local flora and especially have great understanding
of the potential usages of plants (Figueiredo & Smith
2012; Kissanga 2016). In contrast, scientific exploration and documentation of the vegetation of Angola is
still limited. Early botanists such as Friedrich Welwitsch
visited the country in the middle of the 19th century (Welwitsch 1869). The most influential botanist working in
Angola in the first half of the 20th century was John Gossweiler, who worked in all Angolan provinces and collected
over 14,000 specimens. His collection is considered an
especially important source of information for rare and
endemic species (Figueiredo & Smith 2008). Furthermore, Gossweiler produced the first phytogeographic map
of Angola containing 19 principal vegetation types (Gossweiler & Mendonça 1939). Based on this map and his
own observations, Luís A. Grandvaux Barbosa published
a new phytogeographic map in 1971 containing 32 main
1
Revermann et al. | Woody species of the Cusseque area, south-central Angola
season lasting from November to April. The mean annual
precipitation is 987 mm and the mean annual temperature
is 20.4°C (Weber 2013). The study area harbours a high
pedodiversity. The elevated areas are characterized by
deep and developed slightly loamy Arenosols. The slopes
of the smaller valleys of the tributaries and at the western
side of the Cusseque River show shallow Plinthisols on
granitic bedrock. The soils along the eastern part of the
Cusseque River are characterized by very deep and leached
Arenosols. The centre of the valleys support Histosols with
peat layers exceeding 1 m in depth while at the edges of the
wetlands Gleysols are the common soil type (Gröngröft
et al. 2013a).
The main vegetation types covering south central
Angola are semi-deciduous Miombo woodlands and forests
(Figures 2a and b). These woodlands are interspersed with
open vegetation types locally termed anharas de ongote.
The salient feature of the open vegetation types are
dwarf shrubs with a huge underground woody biomass.
This distinct life form was described by White (1976)
as “geoxylic suffrutex”. In the Cusseque area, geoxylic
suffrutices occur on two different soil types: on deep,
leached sandy soils and on shallow, compact, ferralitic
soils. Accordingly, we will differentiate herein between
“geoxylic grasslands on sandy soils” (Figure 2c) and
“geoxylic grasslands on ferralitic soils” (Figure 2d). The
occurrences of the different vegetation types are governed
by topography: woodlands and forests are confined to
the elevated areas and upper slopes. The mid- and lower
and data on the vegetation and the botanical diversity was
available (Revermann 2016). In this study, we present
results of the vegetation survey carried out at the research
site “Cusseque” in the province of Bié located at the upper
reaches of the Okavango River.
matErIaLS aND mEtHODS
Study site
The Okavango River originates on the Angolan Central
Plateau and terminates in a large inland delta in the
Kalahari Desert in Botswana. Within the TFO project,
detailed studies were carried out at four research sites
representing the different parts of the river basin. The
work presented in this paper was carried out at the
study site Cusseque with an area of 100 km2 (13.6985° S,
017.0382° E). The site is located on the Angolan Central
Plateau in the province of Bié (Figure 1; Wehberg &
Weinzierl 2013). The landscape can be described as a
rolling plain intersected by the Cusseque River and its
many tributaries, which are orientated perpendicular to
each other. Three major landscape units can be identified:
the elevated areas, the sloping areas leading down to the
valley bottoms and the valley floors (Gröngröft et al.
2013b). The mean elevation is 1,575 m above sea level
while the difference in elevation between the valley bottom
of the main river and the surrounding elevated areas is
about 100 m (Gröngröft et al. 2013b). The climate of
the Cusseque area is semi-humid with a pronounced wet
Figure 1. Location of the Okavango Basin in southern Africa and the study site “Cusseque” denoted in red. (Projection: WGS 1984; background: RapidEye
high-resolution satellite imagery, recorded 1 May 2013. We acknowledge the DLR for the provision of the data from the RapidEye Science Archive.)
Check List | www.biotaxa.org/cl
2
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
Figure 2. Landscapes of the Cusseque study area: a) Miombo woodland in the middle of the rainy season, b) Miombo woodland at the end of the rainy
season, c) geoxylic grasslands dominated by Cryptosepalum maraviense at the beginning of the rainy season in October, d) geoxylic grassland on sandy
soils at the beginning of the dry season in May; in the background wetland on the valley bottom and Miombo woodlands.
slopes feature geoxylic grasslands. The woodlands and
geoxylic grasslands are separated by ecotones extending
up to several hundred meters where elements of both
vegetation types co-occur. The valley bottoms support
wetlands dominated by Cyperaceae (Revermann et al.
2013; Schneibel et al. 2013).
to the unique character of the geoxylic grasslands, these
were subject to an additional field study. Therein, data were
collected using 10 m × 10 m plots with two 3.3 m × 3.3 m
subplots situated in diagonally opposite corners (adapted
from Dengler 2009). In every plot all vascular plants
found were recorded and their projected cover estimated
visually. Unknown plants were photographed and voucher
specimens were collected according to botanical methods
outlined by Victor et al. (2004). Voucher specimens were
deposited in the herbarium of the ISCED Huíla (LUBA)
and in the Herbarium Hamburgense (HBG). In addition to
the species recorded on the vegetation plots, species found
elsewhere while working at the study area were added to
the checklist. A high number of vegetation plots and several months of field work carried out by four observers in all
seasons ensured a comprehensive coverage of the woody
species present at the study area.
Data collection
Plot based vegetation surveys were carried out during the
growing season in the years 2011 to 2014 and all information is stored in the Vegetation Database of the Okavango
Basin (GIVD ID: AF-00-009, Revermann et al. 2016). In
order to evenly map all existing vegetation units, sampling
followed a random, stratified design. Based on an image
segmentation algorithm using all bands of a Landsat 7
scene, seven major vegetation units were identified. In
these vegetation units random points were created using
GIS and transferred to a hand-held GPS for localization in
the field. Furthermore, additional vegetation plots were
examined in different successional stages of Miombo forest to analyse successional pathways of the regeneration of
natural vegetation after disturbance by shifting cultivation
(Gonçalves et al., accepted). We used a nested plot design
with a 10 m × 10 m plot located in the centre of a 20 m ×
50 m plot. In total, we sampled 148 vegetation plots. Due
Check List | www.biotaxa.org/cl
taxonomy and plant identification
We followed the taxonomy of the checklist “Plants of Angola – Plantas de Angola” by Figueiredo & Smith (2008). We
are aware of recent changes in the taxonomy, but decided
to conform to the national checklist. For identification,
we consulted the Conspectus Florae Angolensis (Exell &
Mendonça 1937, 1951, 1954, 1955; Exell & Fernandes
3
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
1962, 1966; Exell et al. 1970) when possible, and the flora
of neighbouring countries, especially the Flora Zambeziaca
(Exell & Wild 1960) and the field guide to the Trees and
shrubs of Namibia (Mannheimer & Curtis 2009). Additionally, we consulted herbarium collections at the ISCED
Huíla (LUBA) and the Instituto de Investigação Científica
Tropical (LISC) as well as the on-line database JSTOR Plant
Science (http://plants.jstor.org/). For some specimens, we
consulted experts at Kew Botanical Garden (K).
Permits
Permits for plant collection and transfer of biological
material in Angola for scientific purposes was arranged
based on the framework of Material Transfer Agreements
from Angola, negotiated between the Instituto Superior
de Ciências de Educação da Huíla (ISCED, Huíla), Lubango
and the University of Hamburg (UHH), Germany and
authorized on behalf of the Angolan Government by the
Director for Agriculture, Fisheries and Environment of the
Province of Huíla. All International Conventions to which
Angola is signatory country, such as Convention on International Trade in Endangered Species of Wild Fauna and
Flora (1973), Convention on Biological Diversity (1992),
International Treaty on Plant Genetic Resources for Food
and Agriculture (2004), and all other relevant national and
international instruments concerning biodiversity were
taken into account.
Figure 3. The families of woody plants found at the Cusseque study
site, south-central Angola. Only families with more than two species are
shown. In total 37 families of woody plant species occurred, containing
154 species in 99 genera.
and forests showed the highest woody species richness with
110 species belonging to 32 families. A surprisingly high
number of 33 woody species from 14 families were found in
the geoxylic grasslands. These open vegetation types appear
to be merely grasslands in the late growing season (Figures
2c and d) but in fact harbour a remarkable diversity of woody
species (Table 1). In the wetlands few woody species occurred
and we recorded only two Ficus species sporadically occurring along the margins of the wetlands.
One species, Combretum schumannii Engl., was recorded
but is not listed in the current checklist of Angola
(Figueiredo & Smith 2008).
Data analysis
DISCuSSION
We visualized the number of species per family of all
species belonging to one family using the function ‘pie’ in
the statistical software R (R Development Core Team
2016). Data on the frequency of occurrence of a species, the
habitat and the life form were compiled from vegetation
plot data. We assigned frequency according to the following
categories: very rare (1 or 2 observations), rare (3–5
observations), occasional (6–10 observations), frequent
(11–30 observations), common (>30 observations). We
assigned every species to one or more of the following life
form categories based on field observations and literature:
tree, shrub, liana, dwarf shrub and geoxyle. For geoxyles
we followed the definition proposed by White (1976).
White defined a geoxyle as a dwarf shrub that has closely
related species growing as trees and that exhibits massive
woody underground parts.
The 154 species recorded within the 100 km2 of the Cusseque
study site almost equalled the 166 woody species found by
Monteiro (1970) in the entire province of Bié, an area of
70,314 km2. This does not reflect the quality of the study of
Monteiro but rather illustrates how poorly the region has
been surveyed so far. In fact, the study carried out by Monteiro (1970) is of high quality and stands out as the only
study of its time from Angola basing its analysis on quantitative, plot based data. However, the study was restricted to
woodlands and only larger shrubs and trees were included.
In contrast, we included all vegetation types ranging from
woodlands to geoxylic grasslands and wetlands. Due to
the high sampling intensity and the coverage of all vegetation types our species list can therefore be considered a
comprehensive checklist of the woody plant species of the
Cusseque area. However, it must be noted that this list does
not contain any specimen that could not be identified to
at least genus level; some specimens were lacking fruits or
flowers, preventing further identification. Therefore, the
actual number of woody species may be slightly higher.
Despite the relatively recent publication of the checklist
of vascular plants of Angola, subsequent field surveys in
various parts of the country have resulted in additions to
the checklist (Huntley & Coelho 2011). We provided the
first record in Angola of the herbaceous Asteraceae Schistostephium crataegifolium (DC.) Fenzl ex Harv, during the
rESuLtS
We documented 154 woody species belonging to 99 genera
and 37 families (Table 1). The majority of species belonged to
the family Fabaceae (33), followed by Rubiaceae (22), Euphorbiaceae (11), Proteaceae (10) and Combretaceae (9) (Figure
3). Most of the dominant species belonged to the Fabaceae
and occurred with high frequencies. In contrast, the Rubiaceae, second in species richness, contained species occurring
with low frequencies and were less abundant. The woodlands
Check List | www.biotaxa.org/cl
4
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
Table 1. List of species arranged by family. Voucher specimens have been deposited in the herbaria of Lubango (LUBA) and Hamburg (HBG). In most
cases doublets are stored in both herbaria. The herbarium name in parentheses is the location where the specimen used for identification is deposited.
Frequency was assigned according to the following categories: very rare (1 or 2 observations), rare (3–5 observations), occasional (6–10 observations),
frequent (11–30 observations), common (>30 observations). Life forms of the species were assigned to one or more of the categories: tree, shrub, liana,
dwarf shrub and geoxyle.
Species name
Frequency
Life form
Habitat
Collection number(s)
Anacardiaceae
Ozoroa cf. xylophylla (Engl. &Gilg) R.Fern. & A.Fern.
very rare
shrub
geoxylic grassland (sandy soils)
133057B (HBG)
Ozoroa stenophylla Engl. & Gilg.
frequent
shrub
woodland / grassland (ferralitic
soils)
140123 (LUBA)
Rhus arenaria Torre, A.R.
frequent
dwarf shrub /
geoxyle
geoxylic grassland (ferralitic
soils)
140101 (LUBA)
Rhus exelliana Meikle
frequent
dwarf shrub
Rhus gracilipes Exell
frequent
dwarf shrub
woodland / forest
139227; 132483 (HBG)
135250; 134275 (HBG)
Rhus kirkii Oliv.
frequent
dwarf shrub
woodland / forest
139253 (LUBA)
Anisophylleaceae
Anisophyllea boehmii Engl.
frequent
tree
woodland / forest
134316; 139018; 135297 (HBG)
Anisophyllea quangensis Engl. ex Henriq.
rare
dwarf shrub
geoxylic grassland (sandy soils)
133044; 134116 (HBG); 140109 (LUBA)
Annoncaceae
Annona stenophylla ssp. nana Engl. & Diels
rare
dwarf shrub
woodland / forest
133058; 134218 (HBG); 140065 (LUBA)
Uvaria angolensis Welw. ex Oliv. (Figure 4b)
frequent
shrub
woodland / forest
135323; 134240 (HBG)
Xylopia odoratissima Welw. ex Oiv.
frequent
shrub
woodland / forest
133057A; 134263 (HBG)
Xylopia tomentosa Exell
common
shrub
woodland / forest
135279; 132956; 132986 (HBG); 139177
(LUBA)
common
dwarf shrub /
geoxyle
geoxylic grassland (sandy soils)
140121 (LUBA)
Apocynaceae
Chamaeclitandra henriquesiana (Hallier f.) Pichon
Diplorhynchus condylocarpon (Müll. Arg.) Pichon
frequent
shrub / tree
woodland / forest
135300 (HBG)
Landolphia camptoloba (K.Schum.) Pichon
frequent
liana
woodland / forest
132537 (HBG)
Landolphia gossweileri (Stapf ) Pichon
rare
dwarf shrub
geoxylic grassland (sandy soils)
133048 (HBG)
Strophanthus welwitschii (Baill.) K.Schum.
frequent
liana
woodland / forest
135336; 135378; 134091 (HBG)
Asparagaceae
Asparagus sp. 135286
frequent
shrub
woodland / forest
135286 (HBG)
Asparagus cf. africanus Lam.
very rare
shrub
woodland / forest
134115 (HBG)
occasional
shrub
woodland / forest
132695 (HBG)
Parinari capensis Harv.
frequent
dwarf shrub /
geoxyle
geoxylic grassland (sandy soils)
132664; 132898; 140068 (HBG)
Parinari curatellifolia Planch. ex Benth.
common
tree
woodland / forest
132444 (HBG)
Asteraceae
Helichrysum krausii Sch. Bip
Crysobalanaceae
Combretaceae
Combretum acutifolium Exell
very rare
liana / shrub
woodland / forest
135306 (HBG)
Combretum collinum Fresen.
common
tree
woodland / forest
139176 (HBG)
Combretum elaeagnoides Klotzsch
very rare
tree
woodland / forest
132538 (HBG)
Combretum engleri Schinz
frequent
shrub
woodland / forest
133216 (HBG)
Combretum platypetalum ssp. platypetalum Welw.
ex M.A.Lawson (Figure 4f )
occasional
dwarf shrub
geoxylic grassland (sandy soils)
132639; 134114; 140113 (HBG)
Combretum schumannii Engl.
rare
shrub
woodland / forest
139048 (LUBA)
Combretum zeyheri Sond.
frequent
shrub / tree
woodland / forest
135280; 132510 (HBG)
Pteleopsis anisoptera (Welw.) Engl. & Diels
frequent
shrub / tree
woodland / forest
135365; 134110 (HBG); 139066 (LUBA)
Terminalia brachystemma Welw. ex Hiern
frequent
tree
woodland / forest / grassland
(sandy and ferralitic soils)
132997; 134088; 134131 (HBG)
frequent
dwarf shrub /
geoxyle
geoxylic grassland (sandy soils)
140165 (LUBA)
Dichapetalaceae
Dichapetalum cymosum (Hook.) Engl.
Dipterocarpaceae
Monotes africanus A.DC.
common
tree
woodland / forest
132917; 134160; 134228 (HBG)
Monotes angolensis de Wild.
very rare
tree
woodland / forest
132443 (HBG)
Monotes caloneurus Gilg.
rare
tree
woodland / forest
134820 (HBG)
Monotes dasyanthus Gilg
common
tree
woodland / forest
132907; 132961 (HBG); 139228 (LUBA)
Ebenaceae
Diospyros batocana Hiern
occasional
shrub / tree
woodland / forest
139247 (LUBA)
Diospyros chamaethamnus Dinter ex Mildbr.
frequent
dwarf shrub /
geoxyle
geoxylic grassland (sandy soils)
140179 (LUBA)
Continued
Check List | www.biotaxa.org/cl
5
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
Table 1. Continued.
Species name
Frequency
Life form
Habitat
Collection number(s)
Ebenaceae, continued
Diospyros pseudomespilus ssp. brevicalyx Mildbr.
Diospyros virgata (Gürke) Brenan
Euclea crispa ssp. crispa (Thunb.) Gürke
frequent
occasional
frequent
shrub
shrub
dwarf shrub
woodland / forest
woodland / forest
woodland / forest / geoxylic
grassland
135379 (HBG)
132941 (HBG)
135413 (HBG)
very rare
shrub / tree
woodland / forest
139235 (LUBA)
occasional
frequent
rare
rare
common
common
occasional
frequent
common
shrub / tree
shrub / tree
shrub / tree
dwarf shrub
dwarf shrub
dwarf shrub
tree
dwarf shrub
dwarf shrub /
geoxyle
tree
tree
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
geoxylic grassland (ferralitic
soils)
woodland / forest
woodland / forest
139095 (LUBA)
134099; 134135 (HBG); 139068 (LUBA)
139113 (LUBA)
139256 (LUBA)
139238 (LUBA)
139237 (LUBA)
132555; 134232 (HBG); 139038 (LUBA)
132990; 134185 (HBG)
132490; 134199 (HBG)
132691; 132912; 132998 (HBG)
geoxylic grassland (sandy soils)
140167 (LUBA)
frequent
occasional
frequent
common
frequent
dwarf shrub /
geoxyle
tree
shrub / tree
shrub / tree
shrub
shrub / tree
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
134156; 135318 (HBG); 139223 (LUBA)
139065; 135342; 132967 (HBG)
135360; 139242; 133018 (HBG)
135311 (HBG)
132963; 139128 (LUBA)
common
occasional
common
common
common
rare
rare
rare
common
tree
tree
tree
tree
shrub
dwarf shrub
dwarf shrub
dwarf shrub
tree
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
135298; 139016 (LUBA)
132957; 139255 (LUBA)
132676 (HBG)
132900; 135335 (HBG); 139233 (LUBA)
139121 (LUBA)
139257 (LUBA)
139196 (LUBA)
135304 (HBG); 139023 (LUBA)
common
dwarf shrub
132754; 132825 (HBG)
common
dwarf shrub
Dalbergia nitidula Welw. ex Baker
Dialium englerianum Henriq.
Dolichos sp. 140088
rare
frequent
frequent
Entada arenaria Schinz
Eriosema sp. 133109
very rare
rare
shrub / tree
shrub / tree
dwarf shrub /
geoxyle
dwarf shrub
dwarf shrub
Eriosema sp. 132895
rare
dwarf shrub
Erythrina abyssinica Lam. ex DC.
Erythrophleum africanum (Welw. ex Benth.) Harms
Guibourtia coleosperma (Benth.) J.Léonard
Humularia welwitschii (Taub.) P.A.Duvign.
Indigofera baumiana Harms
Indigofera congesta Welw. ex Baker
Kotschya strobilantha (Welw. ex Baker) Dewit & P. A.
Duvign. var. strobilantha
Mucuna sp. 140052
very rare
common
occasional
common
frequent
occasional
rare
140052 (LUBA)
Pericopsis angolensis (Baker) Meeuwen
Pterocarpus angolensis DC
frequent
occasional
tree
tree
tree
dwarf shrub
shrub
dwarf shrub
dwarf shrub /
geoxyle
dwarf shrub /
geoxyle
shrub / tree
tree
geoxylic grassland (ferralitic
soils)
geoxylic grassland (ferralitic
soils)
woodland / forest
woodland / forest
geoxylic grassland (ferralitic
soils)
geoxylic grassland (sandy soils)
geoxylic grassland (ferralitic
soils)
geoxylic grassland (ferralitic
soils)
giant termite mounds
woodland / forest
woodland / forest
woodland / forest
woodland / forest
woodland / forest
geoxylic grassland (ferralitic
soils)
geoxylic grassland (sandy soils)
woodland / forest
woodland / forest
139181 (LUBA)
-
Ericaceae
Erica benguellensis (Welw. ex Engl.) E.G.H. Oliv.
Euphorbiaceae
Bridelia sp. 139095
Hymenocardia acida Tul.
Maprounea africana Müll. Arg.
Phyllanthus angolensis Müll. Arg.
Phyllanthus sp. 139238
Phyllanthus welwitschianus Müll. Arg.
Pseudolachnostylis maprouneifolia Pax
Sclerocroton oblongifolius (Müll. Arg.) Kruijt & Roebers
Uapaca sp. 134199
Uapaca kirkiana Müll. Arg.
Uapaca nitida var. nitida Müll. Arg.
Fabaceae
Abrus melanospermus ssp. suffruticosus (Boutique)
D.K.Harder
Albizia antunesiana Harms
Albizia gummifera (J.F.Gmel) C.A.Sm.
Baphia bequaertii De Wild.
Bauhinia petersiana Bolle
Bobgunnia madagascariensis (Desv.) J.H.Kirkbr. &
Wiersema
Brachystegia bakeriana Hutch. & Burtt Davy
Brachystegia longifolia Benth.
Brachystegia spiciformis Benth.
Burkea africana Hook.
Copaifera baumiana Harms
Crotalaria amoena Welw. ex Baker
Crotalaria cistoides Welw. ex Baker
Crotalaria florida Welw. ex Baker
Cryptosepalum exfoliatum ssp. pseudotaxus (Baker f.)
P.A.Duvign. & Brenan (Figure 4i)
Cryptosepalum exfoliatum ssp. suffruticans
(P.A.Duvign.) P.A.Duvign. & Bre (Figure 4h)
Cryptosepalum maraviense Oliv. (Figure 4g)
common
common
occasional
frequent
135308B; 135620 (HBG)
139236 (LUBA)
133147; 139034 (LUBA)
140088 (LUBA)
134147 (HBG)
133109 (HBG)
132753; 132895 (HBG)
135333 (HBG)
139054 (LUBA)
139146 (LUBA)
132530 (HBG)
139237 (LUBA)
139141 (LUBA)
Continued
Check List | www.biotaxa.org/cl
6
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
Table 1. Continued.
Species name
Frequency
Life form
Habitat
Collection number(s)
Hypericaceae
Psorospermum febrifugum Spach.
rare
shrub / tree
woodland / forest
139036 (LUBA)
Psorospermum tenuifolium Hook.f.
rare
shrub / tree
woodland / forest
132958 (HBG)
common
shrub
woodland / forest
132968; 133005; 133149 (HBG)
Ixonanthaceae
Phyllocosmus lemaireanus (De Wild. & T.Durand)
T.Durand & H.Durand
Lamicaceae
Alvesia rosmarinifolia Welw.
occasional
shrub
woodland / forest
134776; 132533 (HBG)
Tinnea sp. 133121
frequent
dwarf shrub
geoxylic grassland (ferralitic
soils)
133121 (HBG)
Vitex doniana Sweet
occasional
shrub
woodland / woodland ecotone
132915 (HBG)
Vitex madiensis Oliv.
frequent
shrub
woodland / woodland ecotone
132996 (HBG); 139069 (LUBA)
Melastomataceae
Memecylon flavovirens Baker
frequent
shrub / tree
woodland / forest
132519; 133161 (HBG); 139240 (LUBA)
Warneckea sapinii (De Wild.) Jacq.-Fél. (Figure 4d)
occasional
tree
woodland / forest
135309 (HBG); 139140 (LUBA)
occasional
shrub
woodland / forest
132546; 133000; 133096 (HBG)
Meliaceae
Ekebergia benguelensis Welw. ex C.DC.
Moraceae
Ficus pygmaea Welw. ex Hiern
rare
dwarf shrub
wetland margin
141510 (HBG)
Ficus sp. 141539
rare
dwarf shrub
Wetland margin
141539 (HBG)
rare
dwarfshrub /
geoxyle
geoxylic grassland (sandy soils) 140118 (LUBA)
common
shrub
woodland / forest
Myricaceae
Morella cf. serrata (Lam.) Killick
Myrsinaceae
Myrsine africana L.
134107; 134278 (HBG); 139024 (LUBA)
Myrtaceae
tree
woodland / forest
135813 (HBG)
Syzygium guineense ssp. macrocarpum (Engl.) F.White common
Syzygium guineense ssp. barotsense F.White
occasional
shrub / tree
woodland ecotone
135800; 135796 (HBG)
Syzygium guineense ssp. huillense (Hiern) F.White
frequent
dwarf shrub
geoxylic grassland (sandy soils)
133072; 135614; 135882 (HBG)
Ochna afzelii ssp. mechowiana R.Br. ex Oliv.
rare
dwarf shrub /
geoxyle
woodland / grassland
133128 (HBG)
Ochna arenaria De Wild. & T.Durand (Figure 4e)
frequent
dwarf shrub
woodland, geoxylic grassland
(sandy and ferralitic soils)
132947; 133024 (HBG); 140016 (LUBA)
Ochnaceae
Ochna manikensis De Wild.
frequent
dwarf shrub
geoxylic grassland (sandy soils)
132654; 132803 (HBG)
Ochna pulchra Hook.
common
shrub / tree
woodland / forest
135381; 139064 (LUBA)
Ochna pygmaea Hiern
common
dwarf shrub /
geoxyle
woodland / forest, grassland
(sandy soils)
139239; 140154 (LUBA)
Olacaceae
Jasminum pauciflorum Benth.
rare
liana / shrub
woodland / forest
139238 (LUBA)
Schrebera trichoclada Welw.
rare
shrub / tree
woodland / forest
139189 (LUBA)
rare
dwarf shrub
woodland / forest
139033 (LUBA)
common
shrub
woodland / forest
135299 (HBG); 139242 (LUBA)
rare
shrub / tree
woodland / forest
139208 (LUBA)
occasional
tree
woodland / forest
133017 (HBG)
frequent
shrub
woodland / forest
135322; 133032 (HBG); 139164 (LUBA)
Orobanchaceae
Sopubia karaguensis Oliv.
Passifloraceae
Paropsia brazzaeana Baill.
Picodendraceae
Oldfieldia dactylophylla (Welw. ex Oliv.) J.Léonard
Polygalaceae
Securidaca longepedunculata Fresen
Polygonaceae
Oxygonum fruticosum Dammer ex Milne-Redh.
Proteaceae
Faurea intermedia Engl. & Gilg
occasional
shrub / tree
woodland / forest
132720; 139072 (LUBA)
Faurea rochetiana (A.Rich.) Chiov. ex Pic.Serm.
frequent
tree
woodland / forest / ecotone
135307 (HBG)
Faurea saligna Harv.
occasional
tree
geoxylic grassland (ferralitic
soils)
132549; 132980; 134205 (HBG)
Protea baumii Engl. &Gilg.
occasional
dwarf shrub
woodland / forest
132501; 133019; 134225 (HBG)
Protea gaguedi J.F.Gmel.
frequent
tree
woodland / forest
132918 (LUBA)
Protea angolensis var. divaricarta (Engl. & Gilg.) Beard
rare
dwarf shrub
geoxylic grassland (ferralitic
soils)
134200 (HBG)
Protea micans ssp. trichophylla Welw.
occasional
dwarf shrub
geoxylic grassland (sandy soils)
132607 (HBG); 140096 (LUBA)
Proteaceae, continued
Continued
Check List | www.biotaxa.org/cl
7
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
Table 1. Continued.
Species name
Life form
Habitat
Collection number(s)
Protea petiolaris ssp. petiolaris (Hier) Baker & C.H.Wright frequent
Frequency
tree
woodland / forest
132982 (HBG)
Protea cf. welwitschii Engl.
rare
dwarf shrub
geoxylic grassland
(ferraliticsoils)
132480 (HBG)
Protea sp. 133045
rare
dwarf shrub
geoxylic grassland (sandy soils)
133045 (HBG)
rare
shrub
woodland / forest
133093; 135285 (HBG)
rare
dwarf shrub
geoxylic grassland (on sandy
soils) & woodland ecotone
136003 (HBG)
Rhamnaceae
Ziziphus mucronata Willd.
Rubiaceae
Ancylanthos rubiginosus Desf.
Fadogia cf. chrysantha K.Schum.
very rare
shrub
woodland ecotone
134257 (HBG)
Fadogia cf. triphylla var. triphylla Baker
very rare
shrub
woodland / forest
132987 (HBG); 133081 (HBG)
Fadogia fuchsioides Welw. ex Oliv. (Figure 4c)
occasional
shrub
woodland / forest
132524 (HBG)
Fadogia cf. homblei De Wild.
rare
dwarf shrub /
geoxyle
geoxylic grassland
140114 (LUBA)
Fadogia cf. monticola Robyns
rare
dwarf shrub /
geoxyle
geoxylic grassland
140146 (LUBA)
Fadogia sp. 134097
occasional
shrub
geoxylic grassland (ferralitic
soils), woodland ecotone
132453; 134097; 134167 (HBG)
135338 (HBG)
Gardenia brachythamnus (K.Schum.) Launert
very rare
dwarf shrub
woodland ecotone
Keetia cf. gracilis (Hiern) Bridson
very rare
shrub
woodland / forest
132442; 133148 (HBG)
Keetia venosa (Oliv.) Bridson
rare
shrub
woodland / forest
132534A (HBG)
Leptactina benguellensis (Welw. ex Benth. & Hook.f.)
R.D.Good
rare
dwarf shrub
woodland / forest
135313; 135353; 133153 (HBG)
Leptactina prostrata K.Schum
very rare
dwarf shrub
geoxylic grassland (ferralitic
soils)
134181 (HBG)
Pachystigma pygmaeum (Schltr.) Robyns
frequent
dwarf shrub /
geoxyle
geoxylic grassland (sandy soils)
140138 (LUBA)
Pygmaeothamnus cf. chamaedendrum (Kuntze) Robyns very rare
dwarf shrub
geoxylic grassland (sandy soils)
132723 (HBG)
Pygmaeothamnus sp. 132552
dwarf shrub
woodland / forest
132552 (HBG)
very rare
Pygmaeothamnus zeyheri (Sond.) Robyns
rare
dwarf shrub
geoxylic grassland (sandy soils)
132798; 133033; 134089 (HBG)
Rytigynia orbicularis (K.Schum.) Robyns
frequent
shrub
woodland / forest
132925; 134127 (HBG); 139056 (LUBA)
Tapiphyllum cf. psammophilum (S.Moore) Robyns
very rare
shrub
woodland / forest
134279 (HBG)
Tricalysia angolensis A.Rich. ex DC.
very rare
shrub
woodland / forest
132500; 133012 (HBG)
Tricalysia sp. 134221
rare
shrub
woodland / forest
134221 (HBG)
Tricalysia coriacea ssp. nyassae (Benth.) Hiern
occasional
shrub
woodland / forest
133008; 134095; 134170 (HBG)
Tricalysia sp. 135367
very rare
shrub
woodland / forest
135367 (HBG)
rare
dwarf shrub
woodland / forest
139228 (LUBA)
Santalaceae
Thesium sp. 139228
Sapotaceae
Chrysophyllum bangweolense R.E.Fr.
rare
tree
woodland / forest
135359 (HBG)
Englerophytum magalismontanum (Sond.) T.D.Penn.
common
shrub
woodland / forest
135320 (HBG); 133151 (HBG); 139109
(LUBA)
rare
shrub
woodland / forest
135308A (HBG)
Smilacaceae
Smilax anceps Willd.
Strychnaceae
Strychnos cocculoides Baker
frequent
shrub / tree
woodland / forest
139070 (LUBA)
Strychnos pungens Soler.
common
shrub / tree
woodland / forest
139254 (LUBA)
Strychnos spinosa Lam.
occasional
tree
woodland / forest
135301 (HBG)
field work for this study (Gonçalves et al. 2016). In the
case of Combretum schumannii Engl. literature indicated
that the range of the species might extend to Angola
(Exell & Wild 1960). However, neither this taxon nor
its synonyms were included in the Angolan checklist
(Figueiredo & Smith 2008).
Most of the species occurred either in the geoxylic grasslands or in the woodlands and forests. However, many of the
geoxylic suffrutices have closely related tree species growing
nearby in the woodlands (Figures 4h and 4i). In Africa, the
centre of diversity of geoxylic suffrutices is in the Zambezian phytoregion. In regions with similar environmental
Check List | www.biotaxa.org/cl
conditions, such as the Sudanian phytoregion, there is only
a very limited number of geoxylic species (White 1976).
We found two types of geoxylic grasslands, each harbouring a very distinct species pool with only a small overlap.
There is much debate on the environmental factors driving
the emergence of this distinct life form (Davy 1922; White
1976; Maurin et al. 2014; Finckh et al. 2016). However, the
different species composition of the two types of geoxylic
grasslands found in Cusseque can be clearly attributed to the
contrasting edaphic conditions. The two dominant species
in the “geoxylic grasslands on ferralitic soils” Cryptosepalum
maraviense (Figure 4g) and C. exfoliatum ssp. suffruticans
8
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
Figure 4 Typical plants of the Cusseque area: a) Copaifera baumiana, b) Uvaria angolensis, c) Fadogia fuchsioides, d) Warneckea sapinii, e) Ochna arenaria,
f) Combretum platypetalum ssp. platypetalum, g) Cryptosepalum maraviense, h) Cryptoseplaum exfoliatum ssp. suffruticans, i) Cryptosepalum exfoliatum
ssp. pseudotaxus.
(Figure 4i) belong to the Fabaceae. In contrast, the “geoxylic grasslands on sandy soils” were dominated by various
species of the genus Ochna of the Ochnaceae and Parinari
capensis of the Chrysobalanaceae. The “geoxylic grasslands
on ferralitic soils” have their core distribution on the Angolan Central Plateau and make up 8.5% of the land surface
within the Cubango Basin (Revermann et al. in revision).
The “geoxylic grasslands on sandy soils” have a very limited
distribution within the study site and cover 0.7% of the area
of the Cubango Basin. However, they are more extensive
further east in the Cuito River Basin and in eastern Moxico
Province, where they occur on large sandy, alluvial plains of
the Zambezi Graben, e.g., in Cameia National Park.
LItEraturE CItED
Barbosa, L.A.G. 1970. Carta fitogeográfica de Angola. Luanda: Instituto de Investigação Científica de Angola. 323 pp.
Barbosa, L.A.G. 1971. Phytogeographical map of Angola. Mitteilungen der Botanischen Staatssammlung München 10: 114–115.
http://biodiversitylibrary.org/page/15185175
Davy, B.J. 1922. The suffrutescent habit as an adaptation to environment. Journal of Ecology 10: 211–219. http://www.jstor.org/
stable/2255742
Dengler, J. 2009. A flexible multi-scale approach for standardised
recording of plant species richness patterns. Ecological Indicators
9: 1169–1178. doi: 10.1016/j.ecolind.2009.02.002
Exell, A.W. & Fernandes, A. 1962. Conspectus Florae Angolensis.
Lisbon: Junta de Investigações do Ultramar. Vol. 3, Fasc. 1: 1–187.
Exell, A.W. & A. Fernandes. 1966. Conspectus Florae Angolensis.
Lisbon: Junta de Investigações do Ultramar. Vol. 3, Fasc. 2:
189–408.
Exell, A.W., A. Fernandes & E.J. Mendes. 1970. Conspectus Florae
Angolensis. Lisbon: Junta de Investigações do Ultramar and
Instituto de Investigação Científica de Angola. Vol. 4: 401 pp.
Exell, A.W. & F.A. Mendonça. 1937b. Conspectus Florae Angolensis.
Lisbon: Junta de Investigações Colonias. Vol. 1, Fasc. 1: 1–176.
Exell, A.W. & F.A. Mendonça. 1951. Conspectus Florae Angolensis.
Lisbon: Junta de Investigações do Ultramar. Vol. 1, Fasc. 2:
177–422.
Exell, A.W. & F.A. Mendonça. 1954. Conspectus Florae Angolensis.
Lisbon: Junta de Investigações do Ultramar. Vol. 2, Fasc. 1: 1–152.
Exell, A.W. & F.A. Mendonça. 1955. Conspectus Florae Angolensis.
aCKNOWLEDGEmENtS
Research was funded by the German Federal Ministry of
Education and Research (BMBF) in the context of The Future
Okavango (TFO) project, grant number 01LL0912A. We are
grateful for the support of the staff at Kew Royal Botanical
gardens who aided in the identification, in particular David
J. Goyder and Iain Darbyshire. Furthermore, we thank the
people of the villages Kaololo, Sovi, Cusseque and Calomba
and especially the traditional authorities (Sobas) for their
support of our study.
Check List | www.biotaxa.org/cl
9
Volume 13 | Issue 1 | Article 2030
Revermann et al. | Woody species of the Cusseque area, south-central Angola
ommendations for action. Research project final synthesis report
2010–2015. Hamburg: Universtiy of Hamburg, Biocentre Klein
Flottbek. 190 pp.
R Development Core Team. 2016. R: A language and environment
for statistical computing. https://www.R-project.org/
Revermann, R. 2016. Analysis of vegetation and plant diversity patterns in the Okavango basin at different spatial scales — integration
of field based methods, remote sensing information and ecological
modelling [Dissertation]. Hamburg: University of Hamburg. 295
pp. http://ediss.sub.uni-hamburg.de/volltexte/2016/8156/pdf/
Dissertation.pdf
Revermann, R., A.L. Gomes, F.M. Gonçalves, F. Lages & M.
Finckh. 2013. Cusseque — vegetation. Biodiversity and Ecology
5: 59–63. doi: 10.7809/b-e.00247
Revermann, R., A.L. Gomes, F.M. Gonçalves, J. Wallenfang, T.
Hoche, et al. 2016. Vegetation database of the Okavango Basin.
Phytocoenologia 46: 103–104. doi: 10.1127/phyto/2016/0103
Revermann, R., J. Oldeland, F.M. Gonçalves, J. Luther-Mosebach, A.L. Gomes, et al. [In review]. Dry tropical forests of the
Cubango basin in southern Africa — a first classification and
assessment of their woody species diversity. Phytocoenologia.
Schneibel, A., M. Stellmes, D. Frantz, M. Finckh & R. Revermann. 2013. Cusseque — earth Observation. Biodiversity and
Ecology 5: 55–57. doi: 10.7809/b-e.00246
Steudel, T., H. Göhmann, W.-A- Flügel & J. Helmschrot. 2013.
Assessment of hydrological dynamics in the upper Okavango
River Basins. Biodiversity and Ecology 5: 247–261. doi: 10.7809/
b-e.00279
Victor, J.E., M. Koekemoer, L. Fish, S.J. Smithies & M. Mössmer.
2004. Herbarium essentials: the Southern African Herbarium
user manual. Pretoria: SABONET. 93 pp.
Weber, T. 2013. Cusseque — climate. Biodiversity and Ecology 5:
45–46. doi: 10.7809/b-e.00243
Wehberg, J. & T. Weinzierl. 2013. Okavango Basin — physicogeographical setting. Biodiversity and Ecology 5: 11–13. doi: 10.7809/
b-e.00236
Welwitsch, F. 1869. Sertum Angolense, sive stirpium quarundam
novarum vel minus cognitarum. Transactions of the Linnean
Society 27: 1–94. doi: 10.1111/j.1096-3642.1869.tb00202.x
White, F. 1976. The underground forests of Africa: a preliminary
review. The Gardens’ Bulletin Singapore 11: 57–71.
Lisbon: Junta de Investigações do Ultramar. Vol. 2, Fasc. 2:
153–320.
Exell, A.W. & H. Wild. 1960. Flora Zambesiaca. Glasgow: University
Press Glasgow. 336 pp.
Figueiredo, E. & G.F. Smith. 2008. Plants of Angola — Plantas de
Angola, Strelitzia. Pretoria: South African National Biodiversity
Institute. 279 pp.
Figueiredo, E., G.F. Smith & J. César. 2009. The flora of Angola:
first record of diversity and endemism. Taxon 58: 233–236.
http://www.jstor.org/stable/27756837
Figueiredo, E. & G.F. Smith. 2012. Common names of Angolan
plants. Pretoria: Inhlaba Books. 262 pp.
Finckh, M., R. Revermann & M.P.M. Aidar. 2016. Climate refugees
going underground — a response to Maurin et al. 2014. New Phytologist 209: 904–909. doi: 10.1111/nph.13567
Gonçalves, F.M.P., R. Revermann, A.L. Gomes, M.P.M. Aidar,
M. Finckh & N. Jürgens. [Accepted]. Tree species diversity and
composition of Miombo woodlands in south-central Angola,
a chronosequence of forest recovery after shifting cultivation.
International Journal of Forestry Research.
Gonçalves, F.M., J.J. Tchamba & D.J. Goyder. 2016. Schistostephium
crataegifolium Compositae: Anthemideae, a new generic record.
Bothalia — African Biodiversity and Conservation 46: a209. doi:
10.4102/abc.v46i1.202 9
Gossweiler, J. & F.A. Mendonça. 1939. Carta fitogeográphica de
Angola. Lisbon: República Portuguesa Ministério das Colónias.
242 pp.
Gröngröft, A., J. Luther-Mosebach, L. Landschreiber & A.
Eschenbach. 2013a. Cusseque — soils. Biodiversity and Ecology
5: 51–54. doi: 10.7809/b-e.00245
Gröngröft, A., J. Luther-Mosebach, L. Landschreiber, R. Revermann, M. Finckh & A. Eschenbach. 2013b. Cusseque — landscape.
Biodiversity and Ecology 5: 43–44. doi: 10.7809/b-e.00242
Huntley, B.J. & P. Coelho. 2011. Avaliação Rápida Da Biodiversidade
Da Região Da Lagoa de Carumbo/Rapid Biodiversity Assessment
of the Carumbo Lagoon Area. Luanda: Ministério do ambiente,
República de Angola. 219 pp.
Kissanga Vicente da Silva Firmino, R. 2016. Valorização da flora
de Cusseque e Caiúndo no centro e sul de Angola e avaliação
da biomassa lenhosa utilizada para combustível e construção
[Master’s thesis]. Lisbon: University of Lisbon. 63 pp.
Mannheimer, C. & B.A. Curtis. 2009. Trees and shrubs of Namibia.
Windhoek: Macmillan Education Namibia. 526 pp.
Maurin, O., T.J. Davies, J.E. Burrows, B.H. Daru, K. Yessoufou, et
al. 2014. Savanna fire and the origins of the ‘underground forests’
of Africa. New Phytologist 204: 201–214. doi: 10.1111/nph.12936
Monteiro, R.F.R. 1970. Estudo da flora e da vegetação das florestas
abertas do plantalto do Bié. Luanda: Instituto de Investigação
Científica de Angola. 352 pp.
Pröpper, M., A. Gröngröft, M. Finckh, S. Stirn, V. De Cauwer,
et al. 2015. The Future Okavango — findings, scenarios and rec-
Check List | www.biotaxa.org/cl
authors’ contributions: All authors contributed to the manuscript,
carried out fieldwork and worked on the identification of the
collected specimens. RR wrote the initial draft of the manuscript,
analysed the data and compiled the figures and tables.
received: 19 December 2015
accepted: 7 December 2016
academic editor: Alexander Zizka
10
Volume 13 | Issue 1 | Article 2030