ArticlePDF Available


The Malapa fossil assemblage was likely accumulated as a result of a death trap. Given this, the carnivoran species found there must have lived in proximity, close proximity for the smaller species, to the site, offering the possibility of expanding our interpretation of the habitats available to Australopithecus sediba via pinpoint palaeoenvironmental interpretation. To date, the identified carnivorans are the most abundant identified non-hominin taxa at Malapa, and given their territorial behaviour, are important when interpreting the palaeoecology of the site. The extinct false saber-tooth felid (Dinofelis barlowi) suggests that the presence of closed environments and the ancestral form of modern water mongoose (Atilax mesotes) indicates the presence of water in the vicinity. Canids generally support the presence of open habitats. The first appearance in the fossil record of Vulpes skinneri and Felis nigripes indicates the presence of drier open grassland/scrub. The Malapa carnivorans support widespread shifts in carnivore turnover circa 2.0 Ma in Africa and suggest, together with other lines of evidence, the occurrence of a regional transitioning environment during the time of Au. sediba.
1 23
Palaeobiodiversity and
ISSN 1867-1594
Palaeobio Palaeoenv
DOI 10.1007/s12549-016-0245-0
The carnivore guild circa 1.98 million
years: biodiversity and implications for
the palaeoenvironment at Malapa, South
Brian F.Kuhn, Adam Hartstone-Rose,
Rodrigo S.Lacruz, Andy I.R.Herries,
Lars Werdelin, Marion K.Bamford &
Lee R.Berger
1 23
Your article is protected by copyright and all
rights are held exclusively by Senckenberg
Gesellschaft für Naturforschung and Springer-
Verlag Berlin Heidelberg. This e-offprint is
for personal use only and shall not be self-
archived in electronic repositories. If you wish
to self-archive your article, please use the
accepted manuscript version for posting on
your own website. You may further deposit
the accepted manuscript version in any
repository, provided it is only made publicly
available 12 months after official publication
or later and provided acknowledgement is
given to the original source of publication
and a link is inserted to the published article
on Springer's website. The link must be
accompanied by the following text: "The final
publication is available at”.
The carnivore guild circa 1.98 million years: biodiversity
and implications for the palaeoenvironment at Malapa, South
Brian F. Kuhn
&Adam Hartstone-Rose
&Rodrigo S. Lacruz
&Andy I. R. Herries
Lars Werdelin
&Marion K. Bamford
&Lee R. Berger
Received: 4 August 2015 /Revised: 23 March 2016 /Accepted: 5 July 2016
#Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2016
Abstract The Malapa fossil assemblage was likely accumu-
lated as a result of a death trap. Given this, the carnivoran
species found there must have lived in proximity, close prox-
imity for the smaller species, to the site, offering the possibility
of expanding our interpretation of the habitats available to
Australopithecus sediba via pinpoint palaeoenvironmental in-
terpretation. To date, the identified carnivorans are the most
abundant identified non-hominin taxa at Malapa, and given
their territorial behaviour, are important when interpreting the
palaeoecology of the site. The extinct false saber-tooth felid
(Dinofelis barlowi) suggests that the presence of closed envi-
ronments and the ancestral form of modern water mongoose
(Atilax mesotes) indicates the presence of water in the vicinity.
Canids generally support the presence of open habitats. The
first appearance in the fossil record of Vulpes skinneri and
Felis nigripes indicates the presence of drier open grassland/
scrub. The Malapa carnivorans support widespread shifts in
carnivore turnover circa 2.0 Ma in Africa and suggest, togeth-
er with other lines of evidence, the occurrence of a regional
transitioning environment during the time of Au. sediba.
Keywords Paleoecology .Carnivore .Malapa .
Australopithecus sediba
The Malapa fossil locality, north of Sterkfontein and Swartkrans
and due east of Gladysvale (Fig. 1), has yielded a rich assem-
blage of Australopithecus sediba and non-hominin vertebrate
fauna (Berger et al. 2010;Dirksetal.2010; Kuhn et al. 2011).
ThemosaicnatureofAu. sediba anatomy, deemed a Homo-like
australopith, together with its geological age of 1.98 Ma, sug-
gests that this taxon is a potential candidate to occupy a key
evolutionary place in the Australopithecus-Homo transition
(Kivelletal.2011; Kibii et al. 2011;Zipfeletal.2011;
Pickering et al. 2011a; Berger 2013; Churchill et al. 2013;
Schmid et al. 2013; DeSilva et al. 2013). The associated verte-
brate fauna is also remarkable, representing a taxonomically di-
verse assemblage that includes new species as well as last-and-
first appearances of taxa in the Plio-Pleistocene fossil record. To
date, the identified Carnivora dominate the Malapa assemblage.
The identified bovids, equids, and non-hominin primates are less
abundant and hence the ecological interpretations of Malapa
benefit from assessments of the adaptations and habitats associ-
ated with the carnivoran clade. Here, we review the ecology at
Malapa using the carnivorans as an environmental proxy and
further describe the surrounding palaeoenvironments using data
derived from isotopic analysis and identification of fossilized
plant remains in support of the carnivoran assemblage.
*Brian F. Kuhn
Centre for Anthropological Research (CfAR), University of
Johannesburg, House 10, Humanities Research Village, Bunting
Road Campus, Auckland Park 2092, South Africa
USC School ofMedicine, CBA bldg 1, rm. C-36, 6439Garners Ferry
Road, Columbia, SC 29209, USA
Department of Basic Science and Craniofacial Biology, New York
University College of Dentistry, New York, NY 10010, USA
Australian Archaeomagnetism Laboratory, Department of
Archaeology, Environment and Community Planning, La Trobe
University, Melbourne Campus, Bundoora 3086, VIC, Australia
Department of Palaeozoology, Swedish Museum of Natural History,
Box 50007, SE-10405 Stockholm, Sweden
Evolutionary Studies Institute, University of the Witwatersrand,
WITS, Johannesburg 2050, South Africa
Palaeobio Palaeoenv
DOI 10.1007/s12549-016-0245-0
Author's personal copy
Sedimentological and geochronological data support the hy-
pothesis that Malapa should not be considered a widely spread,
time-averaged deposit (Dirks et al. 2010). These data provide
strong support for the view that Malapa was a likely death trap
with rapid deposition and cementation of the skeletal material
deposited on the cave floor (Dirks et al. 2010). The same authors
have hypothesised that the cave system had vertical openings
with no other egress, thus the scenario of animals falling in and
expiring either on impact or over a short period of time. The
vertebrate assemblage is characterised by fairly complete pieces,
often including articulated post-cranial remains consistent with
limited transport of the vertebrate remains before deposition
(Berger et al. 2010;Dirksetal.2010;Kuhnetal.2011). Thus,
it is reasonable to suggest that the carnivoran taxa lived in rela-
tively close proximity to the cave. The tight temporal window
represented by this assemblage enables us to assess the range of
habitats likely available to Au. sediba. The carnivoran sample
recovered from Malapa to date contains eleven taxa, six of which
are identified to species, three to genus, and two only to family
(Table 1). Of note are the last recorded appearance of Dinofelis
barlowi and the first recorded appearance of the black-footed cat
(Felis nigripes) and the new fox species, Vulpes skinneri
(Hartstone-Rose et al. 2013;Kuhnetal.2011).Thesedataon
the last and first appearances at Malapa support widespread turn-
over in the African carnivore guild close to 2.0 Ma in Africa
(Lewis and Werdelin 2010).
The Malapa carnivores and their environments
Habitat-generalist (eurytopic) taxa found at Malapa include
the brown hyaena, Parahyaena brunnea (Kuhn et al. 2011).
Brown hyaenas today are found from the arid beaches of
Namibia (Wiesel 2006; Kuhn et al. 2008), to the Kalahari
Desert (Owens and Owens 1978; Mills and Mills 1978), to
the environments on the outskirts of Johannesburg and
Pretoria, South Africa (Kuhn 2014). Considering the adapt-
ability of the brown hyaenatoday, its presence is of limited use
in establishing the habitat surrounding Malapa.
Members of the family Felidae identified at Malapa pro-
vide two distinct interpretations of the nearby habitats.
Eurytopic large felids identified at Malapa are the leopard
(Panthera pardus) and a specimen identified simply as a large
felid (Kuhn et al. 2011). Leopards are a highly adaptable spe-
cies, occupying environments as diverse as arid deserts to
rainforests (Skinner and Chimimba 2005; Hunter and Barrett
2011). Therefore, the presence of P. p a r d u s at Malapa is con-
sistent with the presence of either open or closed habitats or
both and is of little use in interpreting the habitat surrounding
Malapa. The presence of Dinofelis barlowi, a taxon characterised
by short distal limb elements, which suggests a grappling, am-
bush hunter, and is suggestive of closed environments (Lewis
1995,1997; Marean and Ehrhardt 1995; Turner and Antón
1997; Werdelin and Lewis 2001; Berger and Lacruz 2003;
Lacruz et al. 2006) within its home range. The brachial index
of the known Dinofelis skeletons portrays an animal with rela-
tively short, robust forelimbs capable of greater force than those
of extant leopards, suggesting a forelimb-driven hunting strategy
similar to that of other machairodont felids, such as Megantereon
and Smilodon (Marean and Ehrhardt 1995;Lewis1997;Turner
and Antón 1997; Werdelin and Lewis 2001). The lack of curso-
rial specialisations, combined with powerful forelimbs, further
suggests adaptations to relatively closed habitats, where
Dinofelis could be a more effective ambush predator (Marean
and Ehrhardt 1995; Lewis 1997; Turner and Antón 1997;
Werdelin and Lewis 2001).
The presence of black-footed cat, Felis nigripes,represents
its first and only appearance in the fossil record to date.
F. nigripes is a South African endemic species with highly
constrained home ranges (males 20 km
,females10 km
Fig. 1 Map showing location of Malapa in relation to other fossil sites located within the Cradle of Humankind (modified from Adams et al. 2010)
Palaeobio Palaeoenv
Author's personal copy
(Macdonald et al. 2010). Their habitat is restricted to semi-
desert, Karoo scrub, dry savannah, and open short-grass hab-
itats, and the animal seeks vegetation cover during the day
(Skinner and Chimimba 2005; Macdonald et al. 2010).
Black-footed cats are highly susceptible to habitat loss
(Silwa et al. 2010) and so are key habitat indicators. Thus,
assuming their behaviour has not radically changed, their
presence at Malapa indicates that the cave itself was located
in close proximity to an arid to semi-arid grassland/scrub at
the time of deposition.
A species referable to Genetta is present at Malapa.
Genetta is a widespread genus, with species inhabiting a va-
riety of environments (Lariviére and Calzada 2001; Skinner
and Chimimba 2005). Both G. genetta and G. tigrina current-
ly occur in the region surrounding Malapa (Skinner and
Chimimba 2005; Kuhn 2014) and inhabit all of the likely
biomes surrounding Malapa during the time of deposition
(Lariviére and Calzada 2001). Thus, the presence of Genetta
sp. at Malapa cannot be used to refine environmental charac-
teristics associated with the assemblage but may suggest that
the habitats surrounding Malapa have changed little over time,
which being grasslands in close association with thick wood-
lands supported by spring-fed water sources.
Three genera of mongoose (Herpestidae) have been found
at Malapa. Two of these may represent species of Cynictis and
Rhynchogale although the available material is insufficient to
ascertain species attribution, while the extinct, Atilax cf.
mesotes, is represented by more diagnostic material (Kuhn
et al. 2011). The extant Cynictis penicillata inhabits semi-
arid to arid grasslands, fynbos heath and open bushland, and
maintains home ranges of 0.11.02 km
(Skinner and
Chimimba 2005;HunterandBarrett2011). The extant
Rhynchogale melleri is known from open savannah, open
woodland, and open bushland, but more recent evidence sug-
gests that it may have a broader tolerance of habitats, includ-
ing open grassland (Skinner and Chimimba 2005;Hunterand
Barrett 2011). The presence of termitaria of the genera
Macrotermes and Hodotermes is important to the distribution
of R. melleri (Skinner and Chimimba 2005; Hunter and
Barrett 2011) as these insect genera are generally found in
open woodland and grassland. The third Malapa herpestid,
Atilax cf. mesotes, is an extinct species whose only modern
congener is A. paludinosus, the marsh mongoose. As the com-
mon name suggests, A. paludinosus is usually found along
rivers, streams, marshes, swamps, and tidal estuaries, but
specimens have been found up to half a kilometre from water
sources (Skinner and Chimimba 2005).
Two canids are known from Malapa. They include a large-
sized form represented by a femoral fragment and the smaller
Vulpes skinneri. The extant African fauna includes two species
of canid of a comparable size to the Malapa femur, Lycaon
pictus and Canis simensis. Fossils of the Lycaon genus have
been previously reported in the Cradle of Humankind (Broom
1948;Hartstone-Roseetal.2010; Werdelin and Peigné 2010),
but the Malapa fragment is not adequate for diagnosis below
the family level. While the Malapa specimen can only be
identified as canid, in Africa, large canids are generally indic-
ative of grassland/savannah or open woodland environments.
The genus Vulpes includes a number of species that show clear
preference for open habitats and semi-arid to arid conditions
and commonly occupy small home ranges. The lone excep-
tion to this is V. vulpes, which has proven to be an extremely
adaptable species (Hunter and Barrett 2011). The most impor-
tant feature that sets V. skinneri apart from all other members
of the genus Vulpes (both extant and extinct) is the lack of the
distal accessory cusp on the P
(Hartstone-Rose et al. 2013).
Tabl e 1 Carnivora species identified at Malapa and their associated environmental niche
Family Taxon Environment
Hyaenidae Parahyaena brunnea Eurytopic but includes grasslands/savannah and semi-arid environments
Felidae Dinofelis barlowi
Heavy tree cover
Panthera pardus Eurytopic but includes grasslands/savannah and semi-arid environments
Felis nigripes Grasslands/savannah, scrub, arid and semi-arid environments
Large Felidae indet. Unknown
Viverridae cf. Genetta sp. Eurytopic but includes grasslands/savannah and semi-arid environments
Herpestidae Atilax cf. mesotes
Modern congener prefers a wet environment
cf. Cynictis Semi arid to arid grasslands, fynbos heath and open bushland
cf. Rhynchogale Open savannah, open woodland, and open bushland
Canidae Vulp es sk inn eri
Grasslands/savannah, scrubland, fynbosarid and semi-arid environments
Larger Canidae indet. In Africa tending toward grassland/savannah open woodland and other arid
and semi-arid environments
See text for discussion of environmental assessments and references used to create them
Palaeobio Palaeoenv
Author's personal copy
Considering that this feature is present in all other Vulpes,
including the oldest African species V. riffautae (de Bonis
et al. 2007), it must be considered a derived feature of
V. skinneri that disqualifies it from the direct ancestry of extant
cape fox, V. chama.However,sinceV. skinneri shares many
traits with V. chama, we hypothesise that they shared a recent
common ancestor and likely inhabited similar environments
characterised by semi-arid to arid habitats (Hunter and Barrett
2011), with relatively small territory sizes up to 32.1 km
(Skinner and Chimimba 2005).
Carnivoran guild level assessment of Malapa
The carnivoran guild of Malapa is taxonomically diverse, in-
cluding members of all the extant terrestrial families of
African carnivores except Mustelidae (Kuhn et al. 2011)a
rare group in the Cradle of Humankind fossil record (Brain
1981) aside from Bolts Farm and Plovers Lake (Werdelin and
Peigné 2010). It is also diverse in feeding ecology and body
size range, including large carnivores previously found in the
region (e.g. D. barlowi at Sterkfontein, Mb 4), and a diverse
assemblage of small species, including small herpestids, vi-
verrids, and two taxa hitherto unknown in the fossil record,
F. nigripes and V. skinneri.
The environmental signature of the Malapa carnivore guild
is mixed featuring grassland, woodland, and water in close
proximity to the cave, following previous suggestions based
upon non-carnivore fauna (Reynolds and Kibii 2011).
Although a direct comparison to other environmental recon-
structions such as Sterkfontein, based upon migratory bovids
and pollen from time averaged deposits, would be remiss as
Malapa is a snapshot of that particular environment at a specific
time.Somespecies,suchasV. skinneri,F. nigripes,andthe
tentatively identified Cynictis, indicate the presence of semi-
arid grasslands and scrub lands in direct association with the
cave. An open habitat may also be inferred from the presence of
the larger canid at the site. On the other hand, the identification
of Atilax suggests standing water within about half a kilometre
of the cave, and the existence of Dinofelis barlowi infers the
occurrence of relatively closed or wooded areas in the vicinity
of the cave. The remaining Malapa carnivores are highly eury-
topic and so less relevant as habitat descriptors. What the spe-
cific carnivores can tell us is that Malapa, at 1.98 Ma, sits in a
space where both riverine wooded areas and the beginnings of
more arid scrublands/grasslands met.
The larger carnivore species represented at Malapa are sim-
ilar to those of the older Sterkfontein Mb. 4 assemblage circa
2.62.0 Ma (Herries et al. 2010), which includes D. barlowi,
P. pardus,andP. brunnea. The younger assemblages of
Swartkrans Mb. 1 at 1.91.8 Ma (Pickering et al. 2011b)and
Kromdraai A at 2.01.5 Ma (Vrba 1982; McKee et al. 1995)
have also recorded P. p a rd u s and P. brunnea,butD. barlowi is
there replaced by D. piveteaui. However, ecomorphological
data does not indicate any substantial differences in habitat ad-
aptations between these Dinofelis species (Werdelin and Lewis
2001). Among the smaller carnivores, Vu lpes and Felidae are
represented at Sterkfontein by V
. chama at Jacovec Cave;
<2.4 Ma (Herries et al. 2013)andCaracal caracal at Mb. 2;
2.62.2 Ma (Herries et al. 2013). In addition, Genetta genetta
has been identified from Jacovec Cave, Sterkfontein. On the
other hand, Atilax mesotes is not known from sites older than
type-site for the species. A broader analysis (Lewis and Werdelin
2010) has suggested that there is increased turnover among
South African carnivores at about 2 Ma, and the Malapa assem-
blage may lie just at the cusp of this event as we see the last
D. barlowi and the first appearance of F. nigripes.
The Au. sediba environment
An analysis of a coprolite (likely from a carnivore) recovered
from the site identified the presence of fossilized woody re-
mains of Podocarpus/Afrocarpus spp. among other plant ma-
terial (Bamford et al. 2010). Podocarpus/Afrocarpus are co-
nifer genera associated with moist forested areas of abundant
rainfall (Bamford et al. 2010). Sediment samples from the
hominin layers indicate C
grasses of the type found in sum-
mer rainfall areas, while the analysis of enamel samples from
Malapa Hominin 1 and Malapa Hominin 2 indicate a purely
diet which includes sedge phytoliths (Henry et al. 2012).
Of the plants represented here, those with edible rhizomes are
mostly perennial and inhabit permanent or seasonal wetlands,
whereas sedges from sites with impeded drainage tend to be
annual and have small roots or rhizomes. One palm phytolith
from the family Arecaceae was also recovered, and these trees
require high water tables. Carbon isotope analyses of both
bovid and rodent remains also indicate C
-based diets
(Henry et al. 2012). These data support the presence of nearby
woodland and grassland in the vicinity of Malapa but also
likely suggest mildly cooler temperatures than at present.
Lower temperatures and perhaps a degree of relative
aridification of Southern Africa were also reported based on
the analysis of pollen in deep-sea cores off the Namibian coast
between 2.2 and 2.0 Ma (Dupont et al. 2005;Dupont2006).
Other sources identified an early Pleistocene climate cooling
with development of a stronger atmospheric circulation circa
2.22.0 Ma (Etourneau et al. 2010), whereas aridification re-
lated to the intensification of the Walker Circulation (east-west
atmospheric circulation along the equatorial belt) was detected
at 1.81.7 Ma (Hopley et al. 2007). These drier/cooler condi-
tions identified during the deposition of Malapa are similar to
those described for the broadly contemporaneous Sterkfontein
M5 deposit but not with the older Sterkfontein M4 (Herries
et al. 2010; Herries and Shaw 2011). However, both
Palaeobio Palaeoenv
Author's personal copy
Sterkfontein M4 and Malapa have recorded the presence of
fossilized wood remains (Bamford 1999; Bamford et al. 2010).
Clearly climatic conditions around the deposition of Malapa
were indicative of a cooling trend. Such drier conditions are
consistent with the appearance at Malapa of V. skinneri and
F. nigripes. The occurrence of obligate grazers such as Equus
and the large alcelaphine Megalotragus (Dirks et al. 2010)con-
firms the presence of open grasslands in the greater region.
The Malapa palaeoenvironment has been previously described as
likely including areas of abundant grass and woody vegetation
(Henry et al. 2012). The carnivore assemblage thus provides
additional evidence for open semi-arid to arid grasslands in direct
association with the cave given the small home ranges of
F. nigripes and Cynictis as well as the presence of V. s k i n n e r i .
The presence of A. mesotes also suggests that within a short
distance, there was a source of water. Thus, the Malapa cave
was likely situated either in open habitat or the transitioning
ecozone between the open grasslands and the more wooded
environments along the edge of the valley previously described
by Dirks et al. (2010), with pools of standing water or marsh
(quite possibly spring-fed systems) in close proximity. Although
thepresenceofD. barlowi likely indicates closed environments,
the putative large home ranges for this large body-sized carnivore
(Werdelin and Lewis 2001) make a determination of the proxim-
ity of this habitat to the cave unclear. Podocarpus/Afrocarpus
provides strong evidence for wooded or forested habitats within
the range of D. barlowi and Au. sediba, confirming another
biome in the region. The functional upper limb and thoracic
anatomy of Au. sediba is indicative of adaptations to tree
climbing associated with closed environment (Churchill et al.
2013;Schmidetal.2013). These data suggest that the full range
of palaeoenvironmental conditions available to Au. sediba were
characterised by wooded areas rich in C
grasses and sedges as
well as open grasslands and standing water nearby. It may thus be
important to consider that the continuous aridification of
Southern Africa possibly post ca. 1.9 Ma was a key evolutionary
constraint in the putative transition from Australopithecus to
Homo and may also be indicative of changes in the carnivore
guild at that time.
Acknowledgements We thank the South African Heritage Resource
agency for the permits to work on the Malapa site and the Nash family for
granting access to the Malapa site and continued support of research on their
reserve. Grateful appreciation goes to Sally Reynolds and Jean-Philippe
Brugal for their reviews and comments, which improved the manuscript.
Research by LW is funded by the Swedish Research Council. Additional
support for MB was provided by PAST (Palaeontological Scientific Trust).
AIRH is supported by Australian Research Council Future Fellowship
Compliance with ethical standards
Funding Please see acknowledgements.
Conflict of interest The authors declare that they have no conflict of
Adams, J. W., Herries, A. I. R., Hemingway, J., Kegley, A. D. T., Kgasi,
L., Hopley, P., Reade, H., Potze, S., & Thackeray, J. F. (2010). Initial
fossil discoveries from Hoogland, a new Pliocene primate-bearing
karstic system in Gauteng Province, South Africa. Journal of Human
Evolution, 59(6), 685691. doi:10.1016/j.jhevol.2010.07.021.
Bamford, M. (1999). Pliocene fossil woods from an early hominid cave
deposit, Sterkfontein, South Africa. South African Journal of
Science, 95(May), 231237.
Bamford, M. K., Neumann, F. H., Pereira, L. M., Scott, L., Dirks, P. H. G.
M., & Berger, L. R. (2010). Botanical remains from a coprolite from
the Pleistocene hominin site of Malapa, Sterkfontein Valley, South
Africa. Palaeontologia Africana, 45,2328.
Berger, L. R. (2013). The mosaic nature of Australopithecus sediba.
Science, 340,163.
Berger, L. R., & Lacruz, R. S. (2003). Preliminary report on the first
excavations at the new fossil site of Motsetse, Gauteng, South
Afrcia. South African Journal of Science, 99(May/June), 279282.
Berger, L. R., de Ruiter, D. J., Churchill, S. E., Schmid, P., Carlson, K. J.,
Dirks, P. H. G. M., & Kibii, J.M. (2010). Australopithecus sediba: A
New Species of Homo-Like Australopith from South Africa.
Science, 328,195204.
Brain, C. K. (1981). The hunters or the hunted? An introduction to
African cave taphonomy. Chicago: University of Chicago Press.
Broom, R. (1948). Some South African Pliocene and Pleistocene mam-
mals. Annals of the Transvaal Museum, 21,4749.
Churchill, S. E., Holliday, T. W., Carlson, K. J., Jashashvili, T., Macias,
M. E., Mathews, S., & Berger, L. R. (2013). The upper limb of
Australopithecus sediba.Science, 340(6129), 16. doi:10.1126
de Bonis, L. D., Peigné, S., Guy, F., Likius, A., Mackaye, H. T., Vignaud,
P., & Brunet, M. (2007). The oldest African fox (Vulpes ruffautae n.
sp., Canidae, Carnivora) recovered in late Miocene deposits of
Djurab desert, Chad. Naturwissenschaften, 94,575580.
DeSilva, J. M., Holt, K. G., Churchill, S. E., Carlson, K. J.,Walker, C. S.,
Zipfel, B., & Berger, L. R. (2013).The lower limb and mechanics of
walking in Australopithecus sediba.Science, 340(6129), 15.
Dirks, P. H. G. M., Kibii, J. M., Kuhn, B. F., Steininger, C. M., Churchill,
S. E., Kramers, J. D., & Berger, L. R. (2010). Geological setting and
age of Australopithecus sediba from Southern Africa. Science, 328,
Dupont, L. M. (2006). Late Pliocene vegetation and climate in Namibia
(southern Africa) derived Palynology of ODP site 1082. Geochemistry,
Geophysics, Geosystems, 7(5), 115.
Dupont, L. M., Donner, B., Vidal, L., Pérez, E. M., & Wefer, G. (2005).
Linking desert evolution and coastal upwelling: Pliocene climate
change in Namibia. Geology, 33(6), 461464.
Etourneau, J., Schneider, R., Blanz, T., & Martinez, P. (2010).
Intensification of the Walker and Hadley atmospheric circulations
during the Pliocene-Pleistocene climatic transition. Earth and
Planetary Science Letters, 297(12), 103110.
Palaeobio Palaeoenv
Author's personal copy
Hartstone-Rose, A., Werdelin, L., de Ruiter, D. J., Berger, L. R., &
Churchill, S. E. (2010). The Plio-Pleistocene ancester of wild dogs,
Lycaon sekowei n. sp. Journal of Paleontology, 84(2), 299308.
Hartstone-Rose, A., Kuhn, B. F., Nalla, S., Werdelin, L., & Berger, L. R.
(2013). A new species of fox from the Australopithecus sediba type
locality, Malapa, South Africa. Transactions of the Royal Society of
South Africa, 68(1), 19. doi:10.1080/0035919x.2012.748698.
Bamford,M.,&Berger,L.(2012).ThedietofAustralopithecus sediba.
Nature, 487(7405), 9093.
/n7405/abs/nature11185.html - supplementary-information.
Herries, A. I. R., & Shaw, J. (2011). Palaeomagnetic analysis of the
Sterkfontein palaeocave deposits: implications for the age of the
hominin fossils and stone tool industries. Journal of Human
Evolution, 60(5), 523539. doi:10.1016/j.jhevol.2010.09.001.
Herries, A. I. R., Hopley, P. J., Adams, J. W., Curnoe, D., & Maslin, M. A.
(2010). Letter to the editor: geochronology and palaeoenvironments
of Southern African hominin-bearing localities-A reply to
Wrangham et al., 2009. BShallow-water habitats as sources of fall-
back foods for hominins^.American Journal of Physical
Anthropology, 143(4), 640646. doi:10.1002/ajpa.21389.
Herries, A. I. R., Pickering, R., Adams, J. W., Curnoe, D., Warr, G.,
Latham, A. G., & Shaw, J. (2013). A multi-disciplinary perspective
on the age of Australopithecus in southern Africa. In K. E. Reed, J.
G. Fleagle, & R. E. Leakey (Eds.), Paleobiology of Australopithecus
(pp. 2140). Dordrecht: Springer.
& Kuykendall, K. L. (2007). High- and low-latitude orbital forcing of
early hominin habitats in South Africa. Earth and Planetary Science
Letters, 256(3), 419432.
Hunter, L., & Barrett, P. (2011). A field guide to the carnivores of the
world. Cape Town: Struik Nature.
D. J., & Berger, L. R. (2011). Apartial pelvis of Australopithecus sediba.
Science, 333, 14071411.
Kivell, T. L., Kibii, J. M., Churchill, S. E., Schmid, P., & Berger, L. R.
(2011). Australopithecus sediba hand demonstrates mosaic evolu-
tion of locomotor and manipulative abilities. Science, 333,1411
Kuhn, B. F. (2014). A preliminary assessment of the carnivore commu-
nity outside Johannesburg, South Africa. SouthAfricanJournalof
Wildlife Research, 44(1), 9598.
Kuhn, B. F., Wiesel, I., & Skinner, J. D. (2008). Diet of brown hyaenas
(Parahyaena brunnea) on the Namibian coast. Transactions of the
Royal Society of South Africa, 63(2), 150159.
Kuhn, B. F., Werdelin, L., Hartstone-Rose, A., Lacruz, R., & Berger, L. R.
(2011). Carnivoran remains from the Malapa hominin site, South
Africa. PLoS ONE, 6(11), 111.
Lacruz, R., Turner, A., & Berger, L. R. (2006). New Dinofelis (Carnivora:
Machairodontinae) remains from Sterkfontein Valley sites and a
taxonomic revision of the genus in southern Africa. Annals of the
Transvaal Museum, 43,89106.
Lariviére, S., & Calzada, J. (2001). Genetta genetta.Mammalian Species.
The American Society of Mammalogists, 680,16.
Lewis, M. E. (1995). Plio/Pleistocene carnivoran guilds: implications for
hominid paleoecology. New York: State University of New York at
Stony Brook (Ph.D.).
Lewis, M. E. (1997). Carnivoran paleoguilds of Africa: implications for
hominid food procurement strategies. Journal of Human Evolution,
Lewis, M. E., & Werdelin, L. (2010). Patterns of evolution in eastern and
southern African Carnivora. Paper presented at the Program and
Abstracts, seventieth annual meeting, Society of Vertebrate
Macdonald, D. W., Loveridge, A. J., & Nowell, K. (2010). Dramatis
personae: an introduction to the wild felids. In D. W. MacDonald
& A. J. Loveridge (Eds.), Biology and conservation of wild felids
(pp. 358). Oxford: Oxford University Press.
Marean, C. W., & Ehrhardt, C. L. (1995). Paleoanthropological and paleo-
ecological implications of the taphonomy of a sabertoothsden.Journal
of Human Evolution, 29, 515547.
McKee, J. K., Thackeray, J. F., & Berger, L. R. (1995). Faunal assem-
blage seriation of southern African pliocene and pleistocene fossil
deposits. American Journal of Physical Anthropology, 96,235250.
Mills, M. G. L., & Mills, M. E. J. (1978). The diet of the Brown Hyaena
Hyaena brunnea in the Southern Kalahari. Koedoe, 21,125149.
Owens, M. J., & Owens, D. (1978). Feeding ecologyand its influence on
social organization in Brown Hyenas (Hyaena brunnea,Thunberg)
of the Central Kalahari Desert. East African Wildlife Journal, 16,
Pickering,R., Dirks, P. H. G. M., Jinnah, Z., de Ruiter, D. J., Churchill, S.
E., Herries, A. I. R., & Berger, L. R. (2011a). Australopithecus
sediba at 1.977 Ma and implications for the origins of the genus
Homo.Science, 333,14211423.
Woodhead, J. D. (2011b). Contemporary flowstone development
links early hominin bearing cave deposits in. South Africa Earth
and Planetary Science Letters, 306(12), 2332.
Reynolds, S. C., & Kibii, J. M. (2011). Sterkfontein at 75: review of
palaeoenvironments, fauna and archaeology from the hominin site
of Sterkfontein (Gauteng Province, South Africa). Palaeontologia
Africana, 46,5998.
Schmid, P., Churchill, S. E., Nalla, S., Weissen, E., Carlson, K. J., de
Ruiter, D. J., & Berger, L. R. (2013). Mosaic morphology in the
thorax of Australopithecus sediba.Science, 340(6129), 15.
Silwa, A., Herbst, M., & Mills, M. G. L. (2010). Black-footed cats (Felis
nigripes) and African wildcats (Felis sylvestris): a comparison of
two small felids from South African arid lands. In D. W.
Macdonald & A. J. Loveridge (Eds.), Biology and conservations
of wild felids (pp. 537558). Oxford: Oxford University Press.
Skinner, J. D., & Chimimba, C. T. (2005). The mammals of the Southern
African subregion. Cambridge: Cambridge University Press.
Turner, A., & Antón, M. (1997). The big cats and their fossil relatives.
New York: Columbia University Press.
Vrba, E. S. (1982).Biostratigraphy and chronology, based on Bovidae, of
southern hominid-associated assemblages: Makapansgat,
Sterkfontein, Taung, Kromdraai, Swartkrans; also Elandsfontein
(Saldanha), Broken Hill (now Kabwe) and Cave of Hearths.Nice:
Paper presented at the Proceedings Congress International de
Paleontologie Humaine.
Werdelin, L., & Lewis, M. E. (2001). A revision of the genus Dinofelis
(Mammalia, Felidae). Zoological Journal of the Linnean Society,
132,147258. doi:10.1006/zjls.2OOO.0260.
Werdelin, L., & Peigné, S. (2010). Carnivora. In L. Werdelin & W. J.
Sanders (Eds.), Cenozoic mammals of Africa (pp. 603657).
Berkeley: University of California Press.
Wiesel, I. (2006). Predatory and foraging behaviour of brown hyenas
(Parahyaena brunnea (Thunberg, 1820)) at Cape Fur Seal
(Arctocephalus pusillus pusillus Schreber, 1776) Colonies.
Hamburg: University of Hamburg (PhD).
Zipfel, B., DeSilva, J. M., Kidd, R. S., Carlson, K. J., Churchill, S. E., &
Berger,L.R.(2011).ThefootandankleofAustralopithecus sediba.
Science, 333(6048), 14171420. doi:10.1126/science.1202703.
Palaeobio Palaeoenv
Author's personal copy
That humans originated from Africa is well-known. However, this is widely regarded as a chance outcome, dependant simply on where our common ancestor shared the land with where the great apes lived. This volume builds on from the 'Out of Africa' theory, and takes the view that it is only in Africa that the evolutionary transitions from a forest-inhabiting frugivore to savanna-dwelling meat-eater could have occurred. This book argues that the ecological circumstances that shaped these transitions are exclusive to Africa. It describes distinctive features of the ecology of Africa, with emphasis on savanna grasslands, and relates them to the evolutionary transitions linking early ape-men to modern humans. It shows how physical features of the continent, especially those derived from plate tectonics, set the foundations. This volume adequately conveys that we are here because of the distinctive features of the ecology of Africa.
The use of small-bodied animals, especially small Carnivora, as indicators of palaeoenvironment, is becoming more pronounced. These smaller taxa are more spatiotemporally constrained, owing to high territoriality, ecosystem sensitivity and physical limitations on their ability to travel large distances within their lifespan. This article describes seven new Viverridae specimens, together with a re-evaluation of a previously published specimen, from the hominin-bearing site of Malapa, located in the Cradle of Humankind World Heritage Site, South Africa. When combined with other palaeoenvironmental indicators, the addition of these eight specimens, consisting of Genetta sp., Genetta genetta, and Genetta cf. tigrina, allows for a more precise reconstruction of the environment surrounding Malapa as well as providing additional insights into possible depositional factors that could have resulted in the accumulation of the Malapa deposit.
The Drimolen Palaeocave System in the ‘Fossil Hominid Sites of South Africa’ UNESCO World Heritage Site is well known for numerous remains of early hominins such as Paranthropus robustus and early Homo. These hominin fossils, along with bone tools and notably diverse accumulation of non-hominin primates and fauna, have all been excavated from the 'Main Quarry' area of the site where extensive lime-mining took place. Here we report the first radiometric age of 1.712 ± 0.269 Ma for hominin bearing deposits associated with the DNH7 Paranthropus robustus cranium in the Main Quarry area of the site, which is consistent with recent biochronological estimates. This age is similar to recent estimates for Swartkrans Member 1 Hanging Remnant (somewhere between 2.3 and 1.8 Ma) which also contains Paranthropus and early Homo. Simultaneously, we integrate the newly radiometrically dated Main Quarry deposits with a new fossil deposit, the Drimolen Makondo, discovered in 2013, that is situated some 50 m up the hill to the west from the Main Quarry. It has experienced only limited disturbance from mining but much more extensive erosion. Preliminary excavations and analysis have revealed that the Makondo infill is older than the Main Quarry, dating to 2.706 ± 0.428 Ma. Its greater age is confirmed by biochronology. The Makondo thus overlap with the suggested end of deposition of Australopithecus bearing Sterkfontein deposits, although it is yet to yield any hominin remains. These new dates for the two Drimolen Palaeocave System deposits indicates that, contrary to prior age estimates, the Drimolen site as a whole records the critical hominin and faunal turnover in South African palaeocommunities that occurred around 2.3–1.7 Ma. Finally, as the Drimolen Makondo represents a rare example of a pre-2 Ma fossil bearing deposit in the Gauteng exposures of the Malmani dolomite, we also integrate our results into the greater South African record of palaeodeposit formation (most of which occur between ∼2.0 and 1.0 Ma). An analysis of the age of palaeocave infillings across the Malmani dolomite suggests that, as is classically the case with karst, the height within the dolomite is broadly correlated to their age, although with some notable exceptions that are likely related to localised geological features. Our analysis also indicates that most caves have undergone some form of secondary karstification related to a younger phase of cave formation, contrasting with models that suggest the cavities all formed at the same time and that infill is related to erosion and the opening up of cave passages. As such, the reason that few pre-2 Ma deposits have been identified in the Gauteng exposures of the Malmani dolomite is probably because these older caves have been eroded away. Identifying such early caves is critical in understanding whether earlier hominins may have once existed in South Africa or if erosion of older deposits (or an absence of speleogenesis at this time) has made such early periods absent from the geological record.
Full-text available
A new Plio-Pleistocene viverrid species is described based on two newly discovered maxillae (KW 10141 and KW 10383) from the recent excavations at the hominin-bearing site of Kromdraai (Gauteng, South Africa). This major site allows us to address the conundrum of Paranthropus and Homo origins in South Africa and presents a highly diverse carnivore spectrum (at least 22 species) including herpestids and viverrids. Civettictis braini nov. sp. is a viverrid species comparable in size to the extant African civet Civettictis civetta (Schreber, 1776). However C. braini nov. sp. differs significantly from the extant species in its dental proportions. Its canine and three premolars (P1-P3) are relatively robust, while its carnassials (P4) and two molars (M1, M2) are extremely reduced. This new species supplements our knowledge on carnivore taxonomic diversity and paleoecology in Southern Africa about 2 millions of years ago.
Full-text available
Fossil woods found in the Australopithecus deposits at Sterkfontein Caves, Gauteng, South Africa, are described. The sediments are dated at 2.6-2.8 million years. The woods have been identified as the liana Dichapetalum cf. mombuttense and the shrub, Anastrabe integerrima. Today there is only one species of Dichapetalum, D. cymosum, in South Africa, so the presence of this typically central African gallery forest liana is evidence that at least refugia of dense, humid forest-type vegetation occurred at Sterkfontein during the Pliocene. Anastrabe integerrima grows today on forest margins along and inland from the east and southeast coasts of South Africa. The presence of this plant in the fossil record implies that rainfall was higher during the Pliocene and that gallery forest occurred in an area where today grasslands predominate. Rarely are fossil hominids and plants found together, so this deposit has potential for palaeoclimatic reconstructions.
Seventy-five years after Robert Broom's discovery of the first adult Australopithecus in 1936, the Sterkfontein Caves (Gauteng Province, South Africa) remains one of the richest and most informative fossil hominin sites in the world. The deposits record hominin and African mammal evolution from roughly 2.6 million years (Ma) until the Upper Pleistocene. Earlier excavation efforts focused on the Member 4 australopithecine-bearing breccia and the Member 5 stone tool-bearing breccias of Oldowan and Early Acheulean age. Ronald J. Clarke's 1997 programme of understanding the cave deposits as a whole led to the discovery of the near-complete StW 573 Australopithecus skeleton in the Member 2 deposit of the Silberberg Grotto, and the exploration of lesser known deposits such as the Jacovec Cavern, Name Chamber and the Lincoln Cave. Our aim is to produce a cogent synthesis of the environments, palaeodietary information, fauna and stone artefacts as recorded in the Sterkfontein sequence. We begin with an overview of the site and early accounts of the interpretations of the site-formation processes, after which we discuss each Member in turn and summarize the various types of evidence published so far. Finally, we review the most pertinent debates about the site, including the ages of Sterkfontein Member 2 and 4, and the types of habitats represented at the site through time.
THE FOSSIL POTENTIAL OF MOTSETSE WAS first noted in 1999. This fossil-bearing breccia deposit is located about 16 km northeast of the Sterkfontein caves in the dolomites of the Eccles Formation. The site has been provisionally divided into Upper, Lower, and Middle deposits. Two short field seasons of excavation have been conducted at the site, focusing on the decalcified areas which have yielded more than 2000 macro-mammal specimens, most of which derived from the Lower deposit. No hominin or other primate remains have yet been recovered. Bovids and carnivores are abundant and equids are represented. Isolated teeth of an extinct felid have been attributed to Dinofelis cf. piveteaui. Preliminary descriptions of the fossil material excavated so far are presented here. Based on the presence of the Dinofelis species and its chronological correlation with East African records, 11,16 a provisional palaeontological age of the excavated material is estimated between c. 1.64 and 1.0 Myr. At this stage, no palaeoenvironmental or taphonomic interpretations of the faunal assemblage are suggested pending increased sample sizes and greater stratigraphic resolution of the site.
This paper presents a review of, and new data concerning, the age of Australopithecus in southern Africa. Current dating suggests that Makapansgat Limeworks is the oldest hominin deposit in southern Africa, with Australopithecus africanus dating to between 3.0 and 2.6 Ma. The Taung Child A. africanus fossil from Taung is most likely penecontemporary with the Makapansgat material between 3.0 and 2.6 Ma. A. africanus from Sterkfontein Member 4 is estimated to date to between 2.6 and 2.0 Ma, with the Sts 5 specimen dating to around 2.0 Ma. The A. africanus deposits from Gladysvale are most likely contemporaneous with the Sterkfontein group with an age between 2.4 and 2.0 Ma. The potential second species of Australopithecus, StW 573 from the Silberberg Grotto at Sterkfontein, is most likely dated to between 2.6 and 2.2 Ma. As such, StW 573 is contemporary with A. africanus fossils from Member 4 and suggest that two contemporary Australopithecus species occurred at Sterkfontein between ~2.6 and 2.0 Ma. Based on the presence of Equus the A. africanus fossils from Jacovec Cavern also likely date to Australopithecus sediba-bearing deposits of Malapa date to 1.98 Ma and suggests that three different species of Australopithecus occur in South Africa between 2.3 and 1.9 Ma. Given these dates, A. africanus represents the oldest southern African hominin species being found in two temporally distinct groups of sites, Makapansgat/Taung and Sterkfontein/Gladysvale, and A. sediba is the youngest species at ~1.98 Ma. However, if StW 53 is also Australopithecus, as some have suggested, then this genus survives to younger than 1.8 Ma in South Africa. Australopithecus thus lasted for a significant period of time in southern Africa after the genus is last seen in eastern Africa (Australopithecus garhi at ~2.5 Ma). This new dating indicates that the South African Australopithecus fossils are younger than previously suggested and are contemporary with the earliest suggested representatives of Homo (~2.3 Ma) and Paranthropus (2.7–2.5 Ma) in eastern Africa.
Carnivores of the World is the first comprehensive field guide to all 245 terrestrial species of true carnivores, from the majestic polar bear and predatory wild cats to the tiny least weasel. This user-friendly illustrated guide features 86 color plates by acclaimed wildlife artist Priscilla Barrett that depict every species and numerous subspecies, as well as about 400 line drawings of skulls and footprints. Detailed species accounts describe key identification features, distribution and habitat, feeding ecology, behavior, social patterns, reproduction and demography, status, threats, lifespan, and mortality. Carnivores of the World includes an introduction that provides a concise overview of taxonomy, conservation, and the distinct families within the order Carnivora. More information: