watson.qxd
Check list of the mammals of Tussen-die-Riviere Provincial Nature
Reserve, Free State Province, South Africa
J.P. WATSON
Watson, J.P. 2006. Check list of the mammals of Tussen-die-Riviere Provincial Nature
Reserve, Free State Province, South Africa. Koedoe 49(1): 111–117. Pretoria. ISSN
0075-6458.
Tussen-die-Riviere Nature Reserve falls within the Eastern Mixed Nama-Karoo Biome.
The surface area is approximately 23 000 ha and it is located in a summer rainfall
region. Sixty six different mammal species were recorded for the reserve over the past
13 years which constitutes 65 % of the indigenous mammals recorded for the Free State
Province. The highest number of species for the rodents and insectivores were record-
ed in riverine habitat but the highest abundance were recorded in rocky habitat and was
constituted by only one species, Aethomys namaquensis.
Key words: check list, distribution, mammals, conservation area, South Africa.
J.P. Watson, Scientific Support Services, Free State Department of Tourism, Environ-
mental and Economic Affairs, Private Bag X20801, Bloemfontein, 9300 Republic of
South Africa. (watsonj @dteea.fs.gov.za).
ISSN 0075-6458 111 Koedoe 49/1 (2006)
Introduction
Species lists of occurrence or inventories of
species on nature reserves, national parks or
specific conservation areas have been pub-
lished extensively in the past (Rautenbach
1976; Nel et al. 1980; Pienaar et al. 1980;
Rautenbach 1982; Lynch 1983; Lynch 1989;
Lynch & Watson 1990; Lynch 1994; Avenant
1997; Avenant 2000a). This refined the
knowledge of the distribution of mammal
species in South Africa. This information
formed part of the reference base when the
status of species and distribution delineations
were determined for the biodiversity of
South Africa. Very few provincial reserves in
the Free State (FS) have published mammal
inventories (Rautenbach 1976; Avenant
1997; Avenant 2000b; Avenant & Watson
2002; Wandrag et al. 2002). This is one of the
reasons why the presence of some species in
the FS were omitted or not properly docu-
mented during the compilation of the recent
Red Data Book of the mammals of South
Africa (Friedman & Daly 2004). Lynch
(1983) conducted a survey of the mammals
in the Free State but never sampled any of the
provincial or national conservation areas dur-
ing this survey. All the effort was focused on
private farms, although sampling did occur
adjacent to some provincial reserves.
The intention of this paper is to publish data
that has been collected over the past 13 years
at Tussen-die-Riviere Nature Reserve (TDR)
in the Free State and hopefully it will con-
tribute to the distribution of information on
mammals in central South Africa.
Study area
Tussen-die-Riviere Nature Reserve is situat-
ed in the southern Free State Province of
South Africa, between the Caledon and
Orange rivers, with the western boundary at
the confluence of the two rivers (26º03'–26º
20'E, 30º25'–30º35'S). The reserve was
established in 1967 and proclaimed as a
nature reserve in 1972 (Van Aarde 1984;
Vrahimis 1991). Altitude varies between
1150–1480 m above sea level. The absolute
maximum and minimum temperatures
recorded are 38.3 ºC and -9.3 ºC respective-
ly, with a mean annual precipitation of
441 mm (Vrahimis 1991).
Tussen-die-Riviere is about 23 000 ha in size,
consisting of dolomite koppies with domi-
watson.qxd 2006/04/17 11:02 PM Page 111
nant tree species being Olea, Rhus, Celtis
and Maytenus. The grass plains consist of
Themeda, Aristida, Cymbopogon, Era-
grostis, Chrysocoma, Lesertia, Osteo-
spermum, Salsola and Nestlera species. The
two major rivers, the Orange and Caledon,
that runs through the reserve are fringed by
species such as Diospyros, Phragmitis, Salix
babalonica, Acacia karoo, Lycium and
Celtis. According to Acocks (1988), it falls
within the False Upper Karoo (vegetation
type 300), but Low & Rebelo (1996) rele-
gates it to Eastern Mixed Nama-Karoo (veg-
etation type 52).
Originally TDR was utilised exclusively as a
hunting venue but was also managed as a
conservation area where a number of Artio-
dactyla and Perissodactyla were introduced
into the reserve over the years. Hunting
formed part of the game management actions
of the reserve. Recently the function of TDR
changed from primarily being a hunting
venue to a conservation area and the man-
agement plan was adjusted accordingly.
Materials and methods
Recording of mammal occurrence data has been con-
ducted since 1992. Small mammal surveys were car-
ried out on various occasions utilising Sherman type
live traps, snap traps (museum specials) and catching
specimens by hand. A general bait, consisting of
peanut butter, rolled oats, sunflower oil and beef
extract (Bovril) was used to bait traps. Traps were
checked and re-baited at sunrise and sunset and the
specimens collected were either released after iden-
tification or in cases where animals were sampled
the specimens were deposited in the mammal collec-
tion of the National Museum in Bloemfontein.
The trap success of rodents and insectivores caught
in traps was calculated following the method
described by Rowe-Rowe & Meester (1982).
The following habitat types were sampled: rocky
ridges, riverine vegetation, grass/ shrub land, and old
buildings. In the case of the latter, traps were set out-
side the buildings next to walls in vegetation.
Macro mist nets were used to collect bats at night.
Likely sites were investigated during the day and
bats were collected by hand or shot with .22 dust
shot.
Sight records of various species were taken during
game drives, night drives with a spotlight, and dur-
ing culling operations. Other signs like quills, tracks,
scats and burrows were also used to determine the
presence of some species on the reserve. Sightings
recorded by the reserve personnel were incorporated
in the species lists.
The nomenclature followed in this manuscript is
according to Bronner et al. (2003).
Results and discussion
According to Hilton-Taylor & Le Roux
(1989) the Nama-Karoo and Succulent-
Karoo are two of the most inadequately pro-
tected biomes in South Africa. The Nama-
Karoo, in which TDR is located, is particu-
larly inadequately protected and of concern
(Gelderblom & Bronner 1995). Tussen-die-
Riviere has grassland elements present
which places it in the transitional zone
between the Nama-Karoo and Grassland bio-
mes. Gelderblom et al. (1995) are of the
opinion that the southern Free State is an
unprotected hotspot with high species rich-
ness, endemic and Red Data species and is a
conservation priority area for carnivores.
Augmentation of the conservation area net-
work in this region will protect the maxi-
mum number of species within a minimum
land surface (Gelderblom et al. 1995).
Apart from the alien invasive fallow deer
Cervus dama (Nature Conservation Ordi-
nance No. 8 of 1969) which entered the
reserve by dispersal from neighbouring
farms, only one species of the 66 recorded
species for TDR, viz., impala Aepyceros
melampus, occurs extralimitly in TDR (Plug
& Badenhorst 2001). This species was origi-
nally introduced for hunting purposes. The
species composition (Table 1) on TDR repre-
sents 65 % of the 101 indigenous mammals
for the Free State (Lynch 1983; Watson
1990a; Watson 1990b; Watson 1998; Avenant
& Watson 2002; Ferreira & Avenant 2003).
The Chacma baboon Papio hamadryas ursi-
nus is shown as being introduced in Table 1,
although this species did occur originally in
the region. Baboons that were used in a
behavioural study by a student were released
Koedoe 49/1 (2006) 112 ISSN 0075-6458
watson.qxd 2006/04/17 11:02 PM Page 112
ISSN 0075-6458 113 Koedoe 49/1 (2006)
into the resident troop at TDR in the early
1970s after completion of the project. The
study animals did not originate from this
region but were animals captured at Loskop
Dam Nature Reserve in Mpumalanga. There
is evidence that two genetically different
groups of baboons occur in South Africa, a
northern and southern group (Friedman &
Daly 2004; Bloomer pers. comm.). The result
is that the troop on TDR might have been
contaminated with animals from the northern
group.
Fallow deer, an exotic or alien invasive
species (Nature Conservation Ordinance No 8
of 1969), occasionally enter TDR from
neighbouring farms. The TDR personnel cull
these animals whenever this species is
encountered. Re-entry from adjacent farms
does, however, occur periodically and it is a
continuous process to remove these animals,
underlining the importance of regulating the
importation of alien species into any area.
Ferreira & Avenant (2003) recorded Otomys
saundersiae or Saunders’ vlei rat, Dendro-
mus melanotis the grey climbing mouse, Tat-
era leucogaster the bushveld gerbil, Mys-
tromys albicaudatus the white tailed rat and
Crocidura cyanea the reddish-grey musk
shrew at the reserve. Four other species, Cro-
cidura fuscomurina the tiny musk shrew
(DD), Desmodillus auricularis the short
tailed gerbil, Ichneumia albicauda the white-
tailed mongoose and Lutra maculicollis the
spotted necked otter (NT) were recorded by
Lynch (1983) in close proximity to reserve
but these species have not to date been
recorded on the reserve. Of these four
species, one is near threatened (NT) and one
is data deficient (DD), resulting in two of the
four species listed in a Red Data category as
defined by Friedman & Daly (2004). It is
important to establish whether these species
occur on the reserve, and management prac-
tices should be of such a nature as to be able
to accommodate these species.
Most of the above-mentioned four species
are not common, occur in low numbers or are
very secretive and difficult to observe or cap-
ture.
Rodentia
The rocky outcrop habitat offered the lowest
variety of species. Only one species,
Aethomys namaquensis, was trapped
although sightings of the insectivores Ele-
phantulus myurus, were made while working
in the veld. None were successfully trapped.
Even with the lowest species variety this
habitat had the highest trap success (Table 2).
Aethomys namaquensis is a common and
abundant species in rocky habitat in the Free
State. High trap success percentages are usu-
ally experienced when trapping in rocky
habitat.
Old buildings gave the second highest trap
success with only two species present, viz.,
Mastomys (sensu lato) and Graphiurus mur-
inus. The Mastomys (sensu lato) was trapped
in only two habitats; the other being riverine
vegetation. This species is a generalist and its
presence is also indicative of habitat distur-
bance. Graphiurus murinus was only cap-
tured next to an old building in the Spes
Bona area of the reserve (Table 2). These ani-
mals are, to a large extent, arboreal and old
buildings have shelter and food plants, such
as Diospyros lyciodes, growing along the
walls. Traps were set for this species where
eaten fruits were detected at the old build-
ings. Factors such as shelter, food and signs
of their presence enhanced the trappabilty of
this species at the old buildings although they
do mostly occur in rocky habitats, or in big
trees with holes in the trunks and branches
(Skinner & Smithers 1990).
Grass and shrub plains had the lowest trap
success. Only two species were recorded,
i.e., Aethomys namaquensis and Saccostomys
campestris (Table 2). The former is usually
confined to rocky areas and the inclusion of
the species in this habitat is due to the fact
that traplines were set too close to rocky
areas and an ecotone effect manifested in this
species being trapped in grass/ shrub plains.
The pouched mouse, Saccostomys campes-
tris, is catholic, occupying a variety of vege-
tation and climate types and is widely dis-
tributed (Skinner & Smithers 1990). Lynch
(1983) shows this species to occur only in the
watson.qxd 2006/04/17 11:02 PM Page 113
Koedoe 49/1 (2006) 114 ISSN 0075-6458
Table 1
Mammals recorded in Tussen-die-Riviere
1 = Re-introduced, 2 = Listed Red Data species (Friedman & Daly 2004). 3 = Exotic.
DD = Data Deficient; NT = Near Threatened; VU = Vulnerable.
Scientific and common names are according to Bronner et al. (2003)
Scientific name Common name Status Observed Signs/ TDR
in surveys personnel
Insectivora
2 Suncus varilla Lesser dwarf shrew DD X
2 Atelerix frontalis South African hedgehog NT X
2 Crocidura cyanea Reddish-grey musk shrew DD
Chiroptera
Rhinolophus clivosus Geoffroy’s horseshoe bat X
Neuromicia capensis Cape serotine bat X
Tadarida aegyptiaca Egyptian free-tailed bat X
Primates
1 Papio hamadryas ursinus Chacma baboon X
Cercopithecus pygerythrus Vervet monkey X
Carnivora
Vulpes chama Cape fox X
Canis mesomelas Black-backed jackal X
Aonyx capensis African clawless otter X
Ictonyx striatus Striped polecat X
2 Poecilogale albinucha Striped weasel DD X
Genetta genetta Small-spotted genet X
Suricata suricatta Suricate X
Cynictis penicillata Yellow mongoose X
Galerella pulverulenta Cape grey mongoose X
Atilax paludinosus Marsh mongoose X
Proteles cristatus Aardwolf X
2 Parahyaena brunnea Brown hyaena NT X
Caracal caracal Caracal X
Felis silvestris African wild cat X
Felis nigripes Black footed cat X
Otocyon megalotis Bat-eared fox X
Panthera pardus Leopard X
Perissodactyla
1 Ceratotherium simum White rhinoceros X
2,1 Diceros bicornis minor Black rhinoceros VU X
1 Equus quagga Plains zebra X
Hyracoidea
Procavia capensis Rock hyrax X
Tubulidentata
Orycteropus afer Aardvark X
Artiodactyla
1 Phacochoerus africanus Common warthog X
1 Connochaetus gnou Black wildebeest X
1 Alcelaphus buselaphus Red hartebeest X
1 Damaliscus pygargus phillipsi Blesbok X
Sylvicapra grimmia Common duiker X
Antidorcas marsupialis Springbok X
Rhaphicerus campestris Steenbok X
1 Aepyceros melampus Impala X
1 Oryx gazella Gemsbok X
1 Syncerus caffer African buffalo X
1 Tragelaphus strepsiceros Greater kudu X
watson.qxd 2006/04/17 11:02 PM Page 114
extreme southern Free State Province and is
of the opinion that they are absent from the
rest of the province. This species has howev-
er been recorded at Sandveld Nature Reserve
in the western Free State (Avenant & Watson
2002).
Next to the drainage lines where riverine
vegetation was present, the highest number
of species (5) were recorded. The two most
common species were generalists, Rhab-
domys pumilio and Mastomys (sensu lato).
These two species usually have wide habitat
requirements and are catholic. The high
species variety is probably due to the greater
number of micro habitats that are available,
more diverse food range and the denser veg-
etation cover in these areas. Otomys irrora-
tus, a specialist and vegetarian, was only col-
lected in the riverine areas as this species
tends to keep to areas with firstly dense veg-
etation cover and secondly, areas associated
with higher moisture which also promotes
plant production (Table 2).
Conclusion
As mentioned above, Tussen-die-Riviere
Nature Reserve has 66 recorded mammal
species present. A number of these species
are endemic to South Africa and some are
listed in Red Data book categories (Fried-
ISSN 0075-6458 115 Koedoe 49/1 (2006)
Table 1 (continued)
Scientific name Common name Status Observed Signs/ TDR
in surveys personnel
1 Tragelaphus oryx Eland X
Redunca fulvorufula Mountain reedbuck X
1 Redunca arundinum Southern (Common) reedbuck X
3 Cervus dama Fallow Deer X
Rodentia
Xerus inauris South African ground squirrel X
Pedetes capensis Springhare X
Hystrix africaeaustralis Cape porcupine X
Cryptomys hottentotus African mole-rat X
Otomys irroratus Vlei rat X
Otomys saundersiae Saunders’ vlei rat
2 Mystromys albicaudatus White-tailed rat EN
Saccostomus campestris Pouched mouse X
Malacothrix typica Large eared mouse X
Dendromus melanotis Gey climbing mouse
Rhabdomys pumilio Four-stripe grass mouse X
Mus minutoides (sensu lato) Pygmy mouse X
Mastomys (sensu lato) Multimammate mouse X
Aethomys namaquensis Namaqua rock mouse X
Grahiurus murinus Woodland dormouse X
Tatera brantsii Highveld gerbil X
Tatera leucogaster Bushveld gerbil
Lagomorpha
Lepus saxatilis Scrub hare X
Lepus capensis Cape hare X
Pronolagus rupestris Smith’s red rock rabbit X
Macroscelidea
Elephantulus myurus Rock elephant shrew X
Species in bold are records from Ferreira & Avenant (2003).
watson.qxd 2006/04/17 11:02 PM Page 115
man & Daly 2004). Tussen-die-Riviere
Nature Reserve falls within the transitional
zone between the Nama-Karoo and Grass-
land biomes and is an important area for the
conservation of South African mammals.
The reserve supports a high species richness,
including some endemics and is pointed out
as a priority area for carnivores (Gelderblom
et al. 1995). Furthermore, the reserve forms
an integral part of the conservation network
in this region. This also supports the recom-
mendation by Gelderblom et al. (1995) that
maximum conservation effort with minimum
land surface can be achieved in this region.
This should be investigated by the various
adjoining provinces and an effort must be
made to increase the surface area of the con-
servation areas in this biome. The formation
of a trans-provincial park, linking all the
nature reserves and having a jointly-man-
aged management plan for the whole area
will also enhance the biodiversity conserva-
tion effort in this biome.
Acknowledgements
Gratitude is extended to the Free State Department
of Tourism, Environmental and Economic Affairs for
the time and funds that were available to collect the
data and publish the results. Further gratitude is
extended to all the reserve personnel at Tussen-die-
Riviere who contributed with observations. The
recommendations of Dr Naas Rautenbach, Savvas
Vrahimis and Dr. Pierre Nel on the manuscript are
appreciated and incorporated in the document.
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>>
/JPEG2000ColorImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/AntiAliasGrayImages false
/DownsampleGrayImages true
/GrayImageDownsampleType /Bicubic
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/GrayImageDepth -1
/GrayImageDownsampleThreshold 1.50000
/EncodeGrayImages true
/GrayImageFilter /DCTEncode
/AutoFilterGrayImages true
/GrayImageAutoFilterStrategy /JPEG
/GrayACSImageDict <<
/QFactor 0.15
/HSamples [1 1 1 1] /VSamples [1 1 1 1]
>>
/GrayImageDict <<
/QFactor 0.15
/HSamples [1 1 1 1] /VSamples [1 1 1 1]
>>
/JPEG2000GrayACSImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/JPEG2000GrayImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/AntiAliasMonoImages false
/DownsampleMonoImages true
/MonoImageDownsampleType /Bicubic
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/MonoImageDepth -1
/MonoImageDownsampleThreshold 1.50000
/EncodeMonoImages true
/MonoImageFilter /CCITTFaxEncode
/MonoImageDict <<
/K -1
>>
/AllowPSXObjects false
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0.00000
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/ENU (Use these settings to create PDF documents with higher image resolution for high quality pre-press printing. The PDF documents can be opened with Acrobat and Reader 5.0 and later. These settings require font embedding.)
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>>
>> setdistillerparams
<<
/HWResolution [2400 2400]
/PageSize [612.000 792.000]
>> setpagedevice