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The Paleobiology
of Australopithecus
123
Contributions from the Fourth Stony Brook
Human Evolution Symposium and Workshop,
Diversity in Australopithecus: Tracking the First Bipeds
September 25?28, 2007
Edited by
Kaye E. Reed
School of Human Evolution and Social Change, Institute of Human Origins,
Arizona State University, Tempe, AZ 85287, USA
John G. Fleagle
Department of Anatomical Sciences, Stony Brook University, Stony Brook, NY
11794, USA
Richard E. Leakey
Department of Anthropology and Turkana Basin Institute, Stony Brook
University, Stony Brook, NY 11794, USA
Editors
Kaye E. Reed
School of Human Evolution and Social Change
Institute of Human Origins
Arizona State University
Tempe, AZ
USA
John G. Fleagle
Department of Anatomical Sciences
Health Sciences Center
Stony Brook University
Stony Brook, NY
USA
Richard E. Leakey
Department of Anthropology and Turkana Basin
Institute
Stony Brook University
Stony Brook, NY
USA
ISSN 1877-9077 ISSN 1877-9085 (electronic)
ISBN 978-94-007-5918-3 ISBN 978-94-007-5919-0 (eBook)
DOI 10.1007/978-94-007-5919-0
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Chapter 4
Reconstructing the Habitats of Australopithecus:
Paleoenvironments, Site Taphonomy, and Faunas
Anna K. Behrensmeyer and Kaye E. Reed
Abstract Hominin paleoecology is reconstructed using
many types of evidence from fossils and their geological
context. This evidence is limited by vagaries of the fossil and
geological record. What questions can be asked regarding
Australopithecus ecology given these limitations? We
address this topic by reviewing the major issues concerning
hominin synecology and taphonomy and discuss methods for
deriving ecological information from fossil assemblages and
their geological context. We provide basic information about
the context of the six Australopithecus species known from 22
collecting sites and review their environment of deposition
and other paleoecological evidence. Using this information
we attempt to answer a series of questions, such as whether we
can determine the habitat preferences of the different species,
and whether more than one Australopithecus species shared
an ecosystem at any given place and time. We conclude that
Australopithecus as a genus was eurytopic because of the
wide range of well-documented habitat reconstructions, but
only Australopithecus afarensis, and possibly Australopithe-
cus anamensis, have enough time range and fossil material to
support the interpretation that these species were eurytopic.
The dietary differences between east and south African
species are intriguing given microwear analyses differentiat-
ing the two groups, although the carbon isotope data are
similar. Further evidence of the ecological context of these
species is needed and should be standardized using an
appropriate scale of evidence (temporal and spatial) for the
desired scale of habitat reconstruction.
Keywords Fauna  Paleoecology  Taphonomy
Introduction
Ecological adaptations of early hominins and how these
changed over time are fundamental to understanding human
evolution. Hominin paleoecology can be reconstructed
through various types of evidence contained in fossils and
their geological context. Basic information about hominin
autecology?diet, locomotion, body size dimorphism,
etc.?can be inferred from their anatomy and the isotope
geochemistry of the fossils themselves. Hominin synecol-
ogy, i.e., reconstructions of population structure and abun-
dance, habitat preferences and associations with other
organisms in natural communities, is more elusive, in large
part because hominins are rare components of most fossil
assemblages. Much effort has been devoted to inferring
hominin habitats based on evidence from associated
organisms (e.g., co-occurrence with arboreal mammals
indicating that they lived in a forest community) and geo-
logical evidence for the physical environments and climatic
conditions. Fewer attempts have been made to assess other
aspects of hominin paleoecology, such as population
structure or abundance relative to other taxa.
What do we want to know about the ecology of Aus-
tralopithecus, and how much of what we would like to
know is actually possible, given the limitations of the
geological and paleontological record? These two questions
provide the framework for this paper, which focuses pri-
marily on synecology and approaches to reconstructing the
habitats in which Australopithecus lived. We review the
major issues regarding hominin paleo-synecology and
taphonomy and discuss methods for distilling ecological
information from fossil assemblages and their geological
context. We draw upon examples from the East African
record showing how researchers address various aspects of
the ecological life and times of Australopithecus, and we
also review current interpretations of paleohabitats at
A. K. Behrensmeyer (&)
National Museum of Natural History, Department of
Paleobiology and Evolution of Terrestrial Ecosystems Program,
Smithsonian Institution, P.O. Box 37012NHB MRC 121,
Washington, DC 20013-7012, USA
e-mail: behrensa@si.edu
K. E. Reed
School of Human Evolution and Social Change, Institute of
Human Origins, Arizona State University, Box 874101
Tempe, AZ 85287-4101, USA
e-mail: kreed@asu.edu
K. E. Reed, J. G. Fleagle, R. E. Leakey (eds.), The Paleobiology of Australopithecus,
Vertebrate Paleobiology and Paleoanthropology, DOI: 10.1007/978-94-007-5919-0_4,
 Springer Science+Business Media Dordrecht 2013
41
African Australopithecus sites. Using different scales of
information ranging from documentation of paleoecological
features at individual sites to global-scale climate records
provide a secondary theme for this paper.
The study of Australopithecus synecology draws heavily
upon inferred ecological characteristics of animals, partic-
ularly mammals that were preserved with these hominins.
Information on the sedimentary environments of the sites
and taphonomic attributes of these fossil assemblages also
is necessary for credible interpretations of the associated
fauna. This three-component approach can be applied to
habitat reconstructions for particular fossil assemblages and
also to document habitat variation relating to mammalian
turnover patterns and adaptive shifts associated with dif-
ferent types of habitats. An apparent change in faunal
composition through time can be caused by a shift in
depositional environment or a change in taphonomic pro-
cesses that select for or against certain types of organisms
and skeletal parts. If these confounding variables can be
addressed and corrected for, then it is possible to assess
biological processes that caused turnover, such as the dis-
persal of species out of a region or into a region from
elsewhere, by local speciation and extinction events, and
changes in the relative abundance of persistent lineages.
Conversely, long temporal ranges, broad geographic dis-
tributions of species, or stable patterns of relative abun-
dance are useful for identifying the persistence of similar
habitats through time or across the landscape. Examination
of such patterns in the fossil record can lead to testable
hypotheses regarding the interaction of climate change,
local and regional tectonic processes, and the living com-
munities of plants and animals, thereby providing ecologi-
cal information necessary for understanding large-scale
processes driving hominin evolution.
We begin by outlining major questions regarding Aus-
tralopithecus paleoecology that, ideally, we would like to
answer. We then introduce what is known about the Austra-
lopithecus fossil record, and present three major integrated
approaches to inferring hominin habitats?taphonomy, pa-
leoenvironmental (geological) context, and faunas. This is
followed by summaries of current interpretations of Austra-
lopithecus paleoecology and recommendations for future
research to refine and test these interpretations.
Questions About Australopithecus
Paleoecology
1. What was the range of habitats associated with the genus
Australopithecus, and is it possible to discern each
species? preferred habitat? Did this genus initially live in
forests, woodlands, or other types of closed habitats, or
was it adapted to a mix of open and closed habitats from
its beginnings?
2. What were the important limiting ecological variables
(e.g., food, water, shelter, competition with other spe-
cies, predator avoidance, intra-species interactions) for
Australopithecus?
3. Did the habitats occupied by Australopithecus species
vary across different regions? Was there more than one
Australopithecus species sharing an ecosystem at any one
place and time? How did the later species of Australopi-
thecus co-exist with Paranthropus and early Homo?
4. Was Australopithecus a maker and user of stone tools?
Did any Australopithecus species incorporate significant
meat into its diet?
5. Is there evidence of change through time in a habitat
where the same species continued to exist? Did niche
breadth increase or decrease within the genus Austra-
lopithecus as it evolved?
6. How might global or continental-scale climate change
between 4.5 and 2.0 Ma have affected the paleoecology
of Australopithecus? What was happening in the envi-
ronments of southern versus eastern versus central
Africa, and how do these regional variations compare
with later African climate changes associated with
northern hemisphere glaciation?
These questions represent both possible and impossible
goals for what we can expect to learn from the fossil record.
Answers to many of them depend on both autecological and
synecological evidence. Anatomical data, dental microwear,
and isotopic readings from the hominin fossils themselves
address some of the critical questions regarding australopith
autecology?i.e., what these hominins were functionally
capable of (morphology) and what they actually did in
terms of substrate use, resource use, and other behaviors
(microwear, isotopes, etc.). We do not attempt to review the
vast array of such autecological evidence in this paper.
Instead, we focus on geological context, taphonomic anal-
ysis, and associated fauna, which provide evidence for:
(1) the physical environments and vegetation habitats
occupied by Australopithecus, (2) taphonomic processes
that affected their skeletal remains in the transition from
biosphere to lithosphere, and (3) their distribution through
time relative to changes in paleoenvironments and other
organisms. Sampling biases, especially those relating to
differential preservation of species and time-averaging,
limit what we can know about synecology (see examples
below). One of taphonomy?s important contributions is to
indicate what questions can be realistically pursued with the
42 A. K. Behrensmeyer and K. E. Reed
evidence we have, or are likely to have, from multi-disci-
plinary field and laboratory research.
What We Know: The Basics
At present, six species of early Australopithecus have been
named from three sub-continental regions and *22 col-
lecting sites on the African continent (Figs. 4.1, 4.2;
Table 4.1). Remains are relatively abundant in some of
these sites, including Hadar (Ethiopia) and Sterkfontein
(South Africa), fewer but relatively complete in some such
as Malapa (South Africa), and sparse and fragmentary in
many others. In some cases, fragmentary hominin remains
from the currently documented range of Australopithecus,
i.e., between *4.2 and *2.0 Ma, cannot be certainly
identified as belonging to this genus (see Table 4.1). Much
of what we currently know about the site taphonomy and
paleoecology of Australopithecus is based on a sub-sample
of these sites, including the greater Awash Basin (Hadar,
Maka, Asa Issie, Dikika, Woranso-Mille, Bouri), Laetoli,
and the South African cave sites (Makapansgat,
Sterkfontein).
Fig. 4.1 Map of Africa showing regions and sites in Table 4.1
4 The Habitats of Australopithecus 43
Documented Depositional Contexts for
Australopithecus
? Volcaniclastic plains and paleosols (Laetoli)
? Fluvial channels and floodplains (Lothagam, Kanapoi,
East and West Turkana, Omo Shungura Formation, Ha-
dar, Dikika, Middle Awash)
? Lake margins (East and West Turkana, Hadar, Middle
Awash, Chad)
? Karst terrain and cave deposits (Makapansgat, Sterkfon-
tein, Taung, Gladysvale, Malapa).
Scales and Types of Evidence for
Australopithecus Ecology
The evidence from geological and fossil records includes a
wide range of temporal and spatial scales, each of which
can provide different types of information bearing on
hominin paleoecology (see also Table 4.2):
? Footprints preserve an instant in time, evidence for
hominin behavior such as foraging and social behavior,
and ecological characteristics of contemporaneous (i.e.,
within hours to days) flora and fauna.
? Partial skeleton(s) anatomically informative, represents
the life span of an individual, and if associated in a
contemporaneous death assemblage may provide infor-
mation on group structure.
? Excavation (101?4 m2) provides detailed evidence of the
burial environment and circumstances of the hominin and
any contemporaneous associated fauna and flora, usually
within a short period of time-averaging (*101?
103 years).
? Surface assemblage Fragmentary bones and teeth of
single individuals collected from a surface fossil assem-
blage derived from one or more eroding sedimentary
layers; each specimen represents the life and death of a
single individual but the combined (time-averaged) fau-
nal assemblage may represent *102?105 years.
? Locality (e.g., 104?106 m2) general paleoenvironmental
context and associated fauna from a limited area and
stratigraphic thickness.
? Collecting area, stratigraphic member or sub-member
more time and space typically represented in the com-
bined fossil evidence from these entities, e.g., 104?105
years.
Fig. 4.2 Chronostratigraphic ranges of species of the genus Australopithecus (color coded) based on information from published hominin-
bearing deposits. Dashed lines indicate uncertainty in range limit. See Table 4.1 for references
44 A. K. Behrensmeyer and K. E. Reed
Table 4.1 Pliocene sites in Africa with fossils assigned to the genus Australopithecus, including some for which these records are not certain
based on fragmentary remains, or are likely but not yet published
Collecting
area
Sites Country Habitat
interpretation
Lower
age
Upper
age
Taxon References
Northern
Awash
Basin
Hadar Ethiopia Bushland, open
woodland,
wooded
grassland
3.4 2.9 A. afarensis Campisano (2007),
Campisano and Feibel
(2008), Reed (2008)
Northern
Awash
Basin
Dikika Ethiopia Woodland, open
grasslands
3.8 3.4 A. afarensis Alemseged et al. (2006),
Wynn et al. (2006)
Northern
Awash
Basin
Ledi-Geraru Ethiopia Bushland, open
woodland,
wooded
grassland
3.4 2.95 A. afarensis Geraads et al. (2012)
Northern
Awash
Basin
Woranso-
Mille
Ethiopia Mix of riparian
forest, open
woodland,
grassland
3.8 3.57 A. anamensis, A.
afarensis
Haile-Selassie et al.
(2010a, b)
Middle
Awash
Basin
Asa Issie Ethiopia Closed to grassy
woodland
4.2 4.1 A. anamensis White et al. (2006)
Middle
Awash
Basin
Aramis Ethiopia Grassy woodland
savanna
4.2 4.1 A. anamensis White et al. (2006)
Middle
Awash
Basin
Maka Ethiopia Woodland-
bushland
3.78 3.42 A. afarensis White et al. (1993)
Middle
Awash
Basin
Bouri Ethiopia Lake margin with
grasslands
2.52 2.1? A. garhi Asfaw et al. (1999)
Middle
Awash
Basin
Belohdelie Ethiopia No information 3.7? 3.7? A. afarensis Asfaw (1987)
Southern
Awash
Basin
Galili Ethiopia Woodland to
bushland
4.5 3.5 A. anamensis, A.
afarensis
Kullmer et al. (2008)
Turkana
Basin
Fejej S. Ethiopia No information 4.2? 4.06 A. afarensis? Kappelman et al. (1996)
Turkana
Basin
East Turkana Kenya Riparian forest, wet
grassland,
woodland
4.3 2.7 A. afarensis Kimbel (1988), Brown
et al. (2013)
Turkana
Basin
East Turkana
? Allia Bay
Kenya Mosaic of closed
woodland
and open
grasslands
4.1 3.8 A. anamensis Macho et al. (2003),
Schoeninger et al.
(2003)
Turkana
Basin
West Turkana Kenya Woodland and
forest- edge;
riparian
woodland
4.3 2.5 A. afarensis,
Kenyanthropus
platyops,
Australopithecus sp.
Brown et al. (2013),
Leakey et al. (2001)
Turkana
Basin
Lothagam Kenya Mix of riparian
forest, open
woodlands,
grassland
6.5 5.5 A. afarensis? Hill et al. (1992),
McDougall and Feibel
(1999)
Turkana
Basin
Lothagam Kenya Open, seasonally
dry
3.5 3.5 A. afarensis Leakey and Walker
(2003)
(continued)
4 The Habitats of Australopithecus 45
? Basin a tectonic depression that has accumulated a thick
sequence of sedimentary deposits, representing 105?106
years and providing information on environmental and
paleontological change through time for one sub-region.
? Region tectonic and latitudinal context, comparisons of
different hominin-bearing (and non-hominin-bearing)
habitats through time or across space.
? Continent range of environments, latitudes, habitats, first
and last appearances of hominin species.
? Global climate variation over space and trends and/or
cycles through time.
Paleoenvironments, Taphonomic Biases
and Research Strategies
The paleontological record is imperfect, and taphonomy
often has to provide ??reality checks?? on assumptions about
the biological fidelity of this record and what we can and
cannot know about the past. For australopiths, these limi-
tations may result from the following potential sources of
bias:
1. Small samples of fragmentary remains for any given
hominin taxon may not represent the average or modal
characteristics of that taxon.
2. Even in large samples, selective preservation of hominin
population sub-samples, such as robust individuals and/
or body parts, could skew the range of body sizes and
anatomical features that are available for collection and
study relative to the once-living populations.
3. Available samples of depositional and paleogeographic
contexts where fossil remains of this large-sized primate
occur are likely only partially representative of the range
of habitats and geographic areas where it actually lived.
4. Available assemblages of associated fossil mammals and
hominins represent different degrees of time-averaging
and spatial sampling from the original ecosystems. This
blurs the meaning of ??paleocommunity?? and may bias
comparisons of diversity and other ecological properties
in faunas from different areas, depositional settings, and
time periods (including comparisons to modern faunas).
5. Ecological indicator species may be unevenly preserved
in the fossil record or are difficult to interpret in terms of
their ecological requirements, either due to lack of
modern analogues or to missing body parts.
Table 4.1 (continued)
Collecting
area
Sites Country Habitat
interpretation
Lower
age
Upper
age
Taxon References
Turkana
Basin
Kanapoi Kenya Mix of wooded
and open
grassland
4.17 4.07 A. anamensis Harris et al. (2003)
Turkana
Basin
Omo
(Shungura,
Usno)
Ethiopia Riparian forest
and woodland
3.44 2.44 A. afarensis? A.
garhi?
Brown et al. (2013),
Suwa et al. (1996),
White (2002)
Laetoli Laetolil Fm. Tanzania Mosaic of
woodland,
shrub-
Land, bushland,
grassland
3.8 3.5 A. afarensis Harris et al. (1987), Su
and Harrison (2008),
Kovarovic and
Andrews (2007)
Bahr el
Ghazal
Chad Open grassland
and lake margin
3.5 3 A. bahrelghazali Brunet et al. (1996)
Cave Sterkfontein South
Africa
Open woodland,
riparian forest,
bushland
2.8 2.2 A. africanus, A. sp? Clarke (2013),
Herries et al. (2013)
Cave Makapansgat South
Africa
Mosaic of riparian
woodland,
bushland,
edaphic
grassland
3.5 ? A. africanus Dart (1952), Reed
(1997), Herries et al.
(2013)
Cave Taung South
Africa
Dense woodland 3 2.0? A. africanus Dart (1925), Berger
and Clarke (1995)
Cave Gladysvale South
Africa
Closed/open
vegetation
2.5 1.9 A. africanus Berger and Tobias
(1994)
Cave Malapa South
Africa
No information 2.1 *1.9 A. sediba Berger et al. (2010),
Dirks et al. (2010)
46 A. K. Behrensmeyer and K. E. Reed
We can address the problems above with taphonomic and
paleoenvironmental data in a variety of ways. Obviously,
more data collecting and the opening up of new areas will help
with points (1) and (3), though there will never be enough
fossils to resolve many finer-scale questions about regional
variation and hominin occupation of areas lacking a paleon-
tological record (i.e., most of the African continent). Under-
standing the limitations of the samples that we have, however,
is a big step toward learning how effectively to tackle the
questions that can be answered with the data in hand. There
are ways to calibrate the degree of bias in the preservation of
different body parts, body sizes, and taxa in order to address
Point (2) above. An ??isotaphonomic?? approach that com-
pares samples from specific, well-documented paleoenvi-
ronmental contexts such as fluvial channel lags or lake margin
paleosols can help to control for ecological and taphonomic
variables that differ across environments (Points (3) and (4)).
The use of ??taphonomic control?? taxa, i.e., species with body
Table 4.2 Types of evidence relating the paleoecology of Australopithecus, at increasing spatial and temporal scales, with examples of
autecological and synecological data that can be inferred from this evidence
Evidence Temporal
scale
Spatial scale Examples Autecology of
hominins
Synecology of hominins
Trackways Seconds,
minutes
100?101 m Laetoli footprint
layers
Presence in specific
habitat, on a specific
substrate, behavioral
information
Contemporaneous fauna,
within hours to days
Single individual
with associated
skeletal parts
Lifetime of
the
individual
Habitat of the
individual
Sterkfontein ??Little
Foot??, Hadar
??Lucy??, Dikika
??Salem??
Taxonomic,
ecomorphic,
isotopic information
on body size, diet,
locomotion, etc.;
burial environment
Taphonomic evidence of
scavengers, trauma in
life (e.g., damage to
teeth)
Multiple associated
individuals of a
single taxon
Combined life
span of
individuals
in the
group
Habitat of group Hadar ??First Family??,
South African
Malapa site(?)
Sexual dimorphism,
demography, body
size, diet,
locomotion, burial
environment and
circumstances
Taphonomic evidence of
scavengers, trauma in
life (e.g., damage to
teeth)
Single or multiple
hominin specimens
from a locality,
collecting area, or
well-defined
stratigraphic
interval
103?105 years Habitat area
sampled by
organic
remains, e.g.,
102?105 km2
South African cave
sites, East Turkana,
West Turkana,
Lothagam,
Kanapoi, Hadar,
Bouri, Chad,
Laetoli, etc.
Habitat based on
ecomorphology of
associated fauna
and/or co-
occurrence with
specific ecological
indicator taxa
Community structure and
ecological preferences
inferred from co-
occurring vertebrate taxa
Combined sample
from a geological
formation and
region
105?106 years Area covered by
fossiliferous
deposits and
their source
areas, e.g.,
104?106 km2
Hadar, Middle
Awash, Omo, East
and West Turkana,
Lothagam,
Kanapoi,
Sterkfontein,
Makapansgat
Persistence,
abundance,
disappearance of
individual hominin
taxa through a
stratigraphic interval
Through-time patterns of
mammalian taxonomic
richness, major group
dominance, evenness,
relationships to
environmental
parameters, evidence for
immigration events
Basin with a thick,
partially
continuous
stratigraphic record
105?106 years Basin-scale Turkana Basin,
Awash Basin (Afar
Depression)
Depositional context,
taphonomy, and
ecomorphology of
hominin specimens
within a single basin
through time
Variation in time and space
of faunas and
paleocommunities,
correlation with shifting
physical environments
Region with multiple
localities and
sequences
105?106 years Sub-continental
scale
East Africa, South
African Cave Sites
Variation in
depositional
context, taphonomy,
and ecomorphology
of hominins among
regions
Variation in mammalian
diversity and community
structure in different
tectonic settings,
latitudes, climatic zones
4 The Habitats of Australopithecus 47
size and morphology similar to hominins, such as baboons,
can help to identify variations in abundance of species that are
more likely to be biologically meaningful rather than tapho-
nomically altered (Point (5)).
The ??taphonomic control?? approach was used to com-
pare similar-age portions (Sidi Hakoma (SHT) and Tulu Bor
tuffs) of the Hadar and East Turkana sequences (Behrens-
meyer et al. 2004). Australopithecus is common at Hadar
and rare at East Turkana, but is this the effect of a smaller
fossil sample at East Turkana or a bias against primate
preservation in this area? In both areas, the extinct baboon
Theropithecus and Australopithecus co-occur through the
3.4?2.8 Ma time interval. Similar controlled survey fossil
samples from these two areas indicate that, relative to the
number of specimens of Theropithecus and other large
monkeys recorded in the Hadar Formation and the Tulu Bor
Member of the Koobi Fora Formation, there should be 2.5
hominin specimens in the East Turkana sample if hominins
were as common relative to baboons as they are at Hadar
(Behrensmeyer et al. 2004). However, only one hominin (a
tooth fragment) was found in the Tulu Bor Member. This
suggests (but does not prove) that Australopithecus was less
common in the East Turkana region around 3.4 Ma than at
Hadar. More tests of this kind could improve understanding
of taphonomic versus ecological causes of hominin fossil
abundance.
Two Examples of Site-Based Studies
of Australopithecus Habitats
The types of evidence that feed into habitat reconstructions,
as well as the limitations on inferences imposed by the
fossil record, are illustrated in the following two examples
of well-studied Australopithecus sites in East Africa.
Kanapoi
The Kanapoi locality in the southwestern Turkana Basin,
Kenya, provides evidence for the paleoecology of Austra-
lopithecus anamensis, primarily from fossils preserved in
fluvial sands and paleosols deposited within a time interval
between 4.17 and 4.07 Ma (Harris et al. 2003). These
deposits lie above and below a lacustrine interval, and the
fauna is a time-averaged sample from two similar alluvial
Fig. 4.3 Scale bar showing the different amounts of time-averaging
implied by the paleosol context of each of the two levels at the
Kanapoi Australopithecus anamensis site (left gray box) and the
combined sample of hominins and associated faunal remains from
both levels (right gray box)
48 A. K. Behrensmeyer and K. E. Reed
land surfaces (paleosols) that may have been formed tens of
thousands of years apart (Fig. 4.3). Ecodiversity analysis of
the faunas indicates that the two levels are only slightly
different in terms of the percentage of terrestrial (ground-
dwelling) mammals and the percentage of fresh grass
grazers, i.e., mammals eating more water-dependent/sea-
sonal wetlands grasses (Harris et al. 2003: Figs. 32 and 33;
Behrensmeyer et al. 2007). The combined fauna is used to
characterize the paleoecology of Kanapoi at the time of A.
anamensis and is interpreted as a closed woodland habitat
based on comparisons with analogue environments using
ecological structure analysis (Reed 1997). Other lines of
evidence suggest the existence of open habitats as well,
based on stable isotopic signals in tooth enamel, possible
non-arboreal monkeys, and micromammals, and character-
istics of the paleosols (Wynn 2000; Manthi 2006). Whether
these different habitat types were associated with each other
across space, representing a persistent mosaic environment,
or changed through the interval of time-averaging cannot be
resolved with these analyses.
The amount of time represented by the Kanapoi faunal
samples is clearly long by modern ecological standards and
could include numerous habitat shifts across the areas of
fossil accumulation. Also, the characteristics of the soils are
superimposed on parent sediment that could represent
ecological circumstances different from those during the
period of pedogenesis. The Kanapoi A. anamensis remains
(Leakey et al. 1995) could have been buried (1) during the
initial sedimentary event(s), (2) during the early stages of
pedogenesis affecting this parent material, or (3) later in the
hundreds to thousands of years represented in the two fos-
siliferous units (Behrensmeyer et al. 2007).
Was A. anamensis associated with closed woodland, more
open areas, or a mix of these habitat types? This is an
important question from the standpoint of hominin evolution
because it would indicate either habitat flexibility or speci-
ficity at *4.2 Ma. In the case of shifting habitats through
time, A. anamensis and other species could be closely tied to
one habitat versus another, but still occur as mixed-habitat
fossil assemblages. In the case of a mosaic of both closed and
open habitats, species would have more opportunities, and
perhaps also more selective pressure, to adapt to a variety of
contemporaneous resources and substrates.
The Kanapoi hominins and associated fauna provide one
of the most age-constrained and carefully documented
examples of paleoecological evidence available at present,
but it is still not possible to discriminate between alternative
habitat models because of the amount of ecological time
represented by the combined faunal sample. Mixed-habitat
faunas do not necessarily mean mixed-habitat adaptations
for the species on the faunal list. To improve temporal
resolution, we need better ways of assessing the relative
probabilities of these alternatives, such as more precise
documentation of the depositional and taphonomic history
of the fossil remains in each of the source paleosols, or
stable isotope data from hominin and associated mammals?
tooth enamel (Levin et al. 2011).
Hadar A.L. 333: Environmental Context
of the ??First Family?? Locality
This example shows how the combination of detailed geo-
logical analysis and information from associated faunas
contribute to reconstructing the context of an important
accumulation of at least 15 Australopithecus afarensis indi-
viduals (W. Kimbel, personal communication). The A.L. 333
locality in the Denen Dora Member of the Hadar Formation is
dated at *3.2 Ma and has produced over 260 surface and
excavated specimens of A. afarensis (Behrensmeyer et al.
2003; Behrensmeyer 2008; Harmon et al. 2003). Most of the
hominin fossils were collected along with other faunal
remains from an area of approximately 40 m 9 80 m
(3200 m2) on steep slopes up to the stratigraphic level of 19
excavated specimens. It has long been assumed that the sur-
face hominin fossils were derived from the same sedimentary
unit as the in situ remains, and that this unit was part of a
distinct, carbonate-rich paleosol (Aronson and Taieb 1981).
Further study has shown that the in situ hominin fossils were
buried prior to the formation of overlying paleosols
(Behrensmeyer 2008).
Preserved bedding structures in the fine-grained, homi-
nin-producing strata provide evidence that the abandoned
channel swale continued to aggrade before sustained ped-
ogenesis. The reconstructed paleodrainage of the DD-2
sandstone is oriented south to north with a trunk channel
*40 m wide and 3?5 m deep connecting a tributary system
south of A.L. 333 to a distributary system to the north,
which likely ended on the deltaic plain associated with the
basin?s depositional center. The burial of the hominin
remains in the upper part of the channel involved fine-
grained deposition indicating low-energy, seasonal flood
events, and there is no sedimentological evidence for a
high-energy, catastrophic flood that caused the demise of
the hominins (Behrensmeyer 2008).
Although there is no direct record of vegetation at the
A.L. 333 site, other than CaCO3 root casts associated with
pedogenesis, palynological research in the lower Denen
Dora Member (DD-1 sub-member) suggests that the
regional habitat prior to DD-2 and A.L.333 was predomi-
nantly a dry grassland (Bonnefille et al. 2004). Researchers
(Aronson and Taieb 1981; Bobe and Eck 2001; Reed 2008)
note that fossils of the genus Kobus (waterbuck) and other
reduncines, which indicate moist substrates with ??fresh
grass?? forage (Reed 1997), are common in the Denen Dora
4 The Habitats of Australopithecus 49
Member. Recent geo-faunal analysis by Campisano (2007;
Campisano and Feibel 2008) indicates paleogeographic
differences in the DD-2 sub-member, with edaphic grass-
lands and marshy conditions to the east and more closed,
bush or woodland habitats to the west in the vicinity of A.L.
333. This agrees with stable isotope analysis of pedogenic
carbonates at the excavation site indicating 30?34 % C4
grassland (Hailemichael 2000), which is a relatively low
proportion of grass compared with Hailemichael?s other
samples from the Denen Dora Member.
The in situ hominin remains at A.L. 333 can be related to
a death?and possibly life?association of multiple hominin
individuals with an abandoned channel swale that crossed
an alluvial plain several kilometers from a paleolake to the
north or northeast. The combined evidence indicates that
both wooded and open grassland habitats were present in
the DD-2 sub-member (Reed 2008), with a gradient from
more closed in the west to more open edaphic grasslands to
the east (Campisano 2007; Campisano and Feibel 2008).
Hominins and other animals may have moved along linear
depressions left by abandoned channels when they ventured
across open savanna environments or used such areas for
foraging and shelter. Therefore, as in the Kanapoi example,
it is difficult to specify either open grassland or more bush
to woodland as a ??preferred?? habitat for the A.L. 333 A.
afarensis; the conservative interpretation is that they were
associated with a mix of these types of vegetation.
Paleoenvironmental context provides only part of the
history of the A.L. 333 hominin assemblage, and ongoing
research is investigating alternative scenarios for the accu-
mulation of the hominins based on taphonomic evidence
from the fossils themselves, their spatial patterns of pres-
ervation, and co-occurring organisms (Behrensmeyer et al.
2003; Harmon et al. 2003). These scenarios cover a range of
temporal scales and processes of death and burial (Fig. 4.4)
and additional taphonomic analysis likely will shed new
light on the paleoecology of this unusual fossil hominin
accumulation.
Using Faunas to Infer Hominin Habitats
Today African habitats range from rain forests to deserts.
The amount of rainfall, temperature, sunlight, evapo-tran-
spiration, soil type, landscape physiography, and weather
patterns/seasonality are the abiotic factors that cause dif-
ferentiation in habitats. Floras and faunas are sensitive
Fig. 4.4 Scale bar showing the different amounts of time-averaging
that would be implied by alternative scenarios for the taphonomic
origin of the A.L.333 A. afarensis assemblage. The biological and
behavioral meaning of this as a population sample depends on which
scenario is supported by paleoenvironmental context and taphonomic
evidence
50 A. K. Behrensmeyer and K. E. Reed
indicators of these environmental conditions, even on a
relatively small spatial scale. Thus, ecological analysis of
fossils provides a window into past habitats, which in turn
can be used to reconstruct climatic conditions (Archibold
1995; Andrews 2006). In the tropical belt, the seasonal
pattern and the amount of rainfall are the most important
determining factors of the vegetation physiognomy (Haw-
kins et al. 2003). Habitats of various types often occur
together in a particular spatial region because of changes in
soil types, subterranean water, etc. For example, it is pos-
sible to have forests along rivers adjacent to near desert-like
habitats, a condition that occurs where the present-day
Awash River flows through the Afar hominin fossil beds in
Ethiopia. These habitats are either called ecotonal or
mosaic. Often mosaic habitats are indicated by ecological
analysis of fossil assemblages; if this is due to time aver-
aging of shifting habitats then the reconstruction of a con-
temporary mosaic of habitats could be incorrect. On the
other hand, varying faunal compositions from time-syn-
chronous collections over a broad spatial area, would lend
support to the interpretation of a mosaic habitat structure.
Occasionally, it is possible to reconstruct the habitat asso-
ciated with hominin remains in a small spatial region and
arrive at an interpretation for a non-mosaic (homogeneous)
habitat at this scale (e.g., White et al. 2009).
Patterns of species occurrence at particular sites and their
persistence and turnover through stratigraphic successions,
combined with ecomorphic features of these species, pro-
vide evidence for ecological characteristics of hominin
species and even for different populations of the same
species (e.g., A. afarensis at Laetoli and Hadar; Su and
Harrison 2008). Regional patterns can be combined in
studies of larger-scale biogeographic and ecological pat-
terns across the African continent. When compared with
independently documented habitat shifts, species turnover
patterns at individual sites may provide information on the
eurytopic (??adaptable??) and stenotopic (??specialized??)
nature of lineages, including hominins. One might expect
that eurytopic species would occur consistently through
time, despite habitat shifts, and across the landscape in a
variety of habitats. In contrast, stenotopic species may only
be recovered if particular habitats are sampled and may be
consistently fewer in fossil assemblages, perhaps suggesting
movement in and out of regions through time in response to
habitat fluctuations. Over time stenotopic lineages may
exhibit higher extinction and diversification rates (Vrba
1980; Badgley et al. 2008).
Because of collection practices, time-averaging, and
spatial restrictions, it is probable that most fauna-based
habitat reconstructions of Pliocene hominin localities rep-
resent a temporal (time-averaged) scale of 104?105 years, as
illustrated in the Kanapoi example above, a relatively
coarse level of resolution that may incorporate numerous
shorter-term ecological shifts. On the other hand, recon-
structions based on paleosols and pollen from specific sites
may signal habitats of small area or short duration that may
or may not be associated with the place and time where the
sampled vertebrate fauna or hominins actually lived.
Paleoecological Evidence and Current
Interpretations of Australopithecus Sites
The following section reviews various Australopithecus
taxa (Table 4.1) and the information that is known about the
paleoecological context of each locality.
Sites with Hominins of Uncertain Taxonomic
Assignment
? Lothagam Hill, Kenya. There is abundant fauna from
Lothagam, but hominins are very rare throughout the
7.0?3.5 Ma time span. Only two teeth are known from
*6.5?5.5 Ma in the upper Nawata Formation and one
poorly preserved mandible from the overlying Apak
Member of the Nachukui Formation (Leakey and Walker
2003). The bovid fauna of the upper Nawata is dominated
by aepycerotins, alcelaphins, and reduncins, indicating a
mix of gallery forest, open woodlands and grasslands.
Fewer alcelaphins and more tragelaphins in the Apak
Member as well as an increase in colobines provide
evidence for a more closed habitat at *5.0 Ma (Leakey
and Harris 2003), although d13C analysis of Apak
Member bovid tooth enamel indicates a significant
component of C4 vegetation (Cerling et al. 2003). The
loss of Etheria (oyster) reefs in the Apak Member indi-
cates a change to an ephemeral flow regime. Carbon
isotope analysis of pedogenic carbonates and tooth
enamel through the Lothagam succession indicates ??a
mosaic ecosystem with stands of pure C3 vegetation
interspersed with mixed C3/C4 floras?? but no pure C4
grasslands (Cerling et al. 2003). Given the number and
excellent preservation of other mammalian fossils, the
scarcity of hominins throughout the Nawata Formation
indicates this group was rare to absent in Lothagam?s late
Miocene paleocommunity (Leakey and Harris 2003).
? Omo (Shungura Formation), Ethiopia. There are thousands
of faunal specimens from this locality, largely consisting of
isolated teeth, including some attributed to Australopithecus
(Suwa et al. 1996) or more recently to Australopithecus
garhi (White et al. 2002). Through the 1.2 Myr of likely
Australopithecus occupation of this environment, the fossils
4 The Habitats of Australopithecus 51
derive from fluvial depositional settings associated with the
paleo-Omo River. The habitats associated with the hominins
include riparian forest and woodland habitats from 3.2 to
2.0 Ma; alcelaphins and antilopins are a notably small
component of the fauna during this time, indicating that open
grassland habitats were limited in extent in the paleo-Omo
River Valley (Bobe and Eck 2001; Bobe et al. 2002;
Alemseged et al. 2007).
? West Turkana, Kenya. A number of hominin remains are
identified as A. afarensis (Leakey et al. 2001), and at least
42 catalogued, but unpublished, specimens are assigned
to Australopithecus (E. Mbua, personal communication).
The fossils are mostly teeth from above the 3.4 Ma Tulu
Bor Tuff, in the Lomekwi Member of the Nachukui
Formation. Kenyanthropus platyops also occurs in the
Lomekwi and underlying Kataboi Member, indicating the
presence of two contemporaneous hominin genera. Based
on the bovid fauna, the habitat of the lower through upper
Lomekwi members has been interpreted as a mosaic
dominated by woodland and forest-edge vegetation
(Harris et al. 1988; Leakey et al. 2001). This is supported
by abundant Theropithecus brumpti, a species regarded as
indicating more closed habitats than T. darti, which is
common in the contemporaneous Hadar Formation in
Ethiopia (Leakey et al. 2001).
A. bahrelghazali
? Bahr el Ghazal, Chad. This site is dated between 3.0 and
3.5 Ma and is the only central African site from which
any Australopithecus species has been recovered. The
fauna associated with this hominin lacks tragelaphins and
aepycerotins but has abundant alcelaphins, reduncins, and
antilopins, indicating open grassland and lake margin
habitats (Geraads et al. 2001).
A. anamensis
? Allia Bay, Kenya. Hominin remains consisting mostly of
isolated teeth are preserved in a fluvial channel lag con-
text associated with the base of the Moiti Member at
*4.0 Ma. Based on analysis of stress lines in the enamel
of fossil herbivore teeth from this channel deposit, Macho
et al. (2003) suggest that the habitat of A. anamensis was
quite seasonal and similar to Masaai Mara in Kenya
today. Schoeninger et al. (2003), using carbon and oxy-
gen stable isotope analysis of tooth enamel, infer a
mosaic habitat of closed woodland and grasslands with
higher rainfall than the region receives today.
? Kanapoi, Kenya. A total of 59 specimens of A. anamensis
have been reported from this locality. The abundant
associated fauna is derived from floodplain paleosols and
distributary sands that span an estimated total time period
of about 100 kyr (see earlier discussion about Kanapoi
time-averaging and habitat reconstruction) (Harris et al.
2003). Faunal eco-diversity analyses of these two levels
are similar and indicate either wooded habitat or a mosaic
with wooded and more open areas, while stable isotopes,
the possible non-arboreal monkeys, and micromammals
indicate presence of open grasslands. Wynn (2000) sug-
gests, based on the characteristics of the paleosols where
hominin remains were recovered in situ, that A. anam-
ensis at least occasionally was associated with open
conditions within a spatially variable ecosystem, typified
by a mosaic of habitats, ranging ??from forb-dominated
edaphic grassland to gallery woodland, providing a larger
view of the mixed ecosystem in which A. anamensis
lived.??
? Aramis and Asa Issie, Ethiopia. White et al. (2006)
recovered A. anamensis from two localities near Aramis
in the Middle Awash. The Asa Issie fauna has high per-
centages of colobine monkeys and tragelaphine bovids as
well as forest-adapted avifauna and micromammals
leading these authors to interpret the habitat as closed to
grassy woodlands. The Aramis A. anamensis locality
lacks other fauna but stable carbon analysis of pedogenic
carbonate provide an average of *25?35 % C4, inter-
preted as indicating a ??humid, grassy, woodland savan-
nah environment.?? (White et al. 2006: 885).
? Woranso-Mille, Ethiopia. Haile-Selassie et al. (2010b)
report a sample of 26 hominin remains of Australopi-
thecus, recovered from the northernmost locality in the
Afar thus far and dated to *3.57?3.8 Ma. These fossils
consist of isolated teeth and partial mandibles and max-
illae that exhibit features of both A. anamensis and A.
afarensis, thus a possible transitional form. The fauna
from four collection sites indicates a mix of riverine
forest, open woodland and grassland habitats, based on
relatively abundant Theropithecus oswaldi aff. darti and
tragelaphin, aepycerotin, and bovin bovids, which Haile-
Selassie et al. (2010a) note is more similar to the older
Kanapoi fauna than that of age-contemporaneous Laetoli
(see below).
? Galili, Ethiopia. This site has produced Australopithecus
teeth and a femur (Kullmer et al. 2008; Viola et al. 2008)
identified as most similar to A. anamensis. The fauna
suggest a comparable date with Kanapoi, and the Kataboi
Member of the Nachukui Formation, although there are
some similar fauna with the younger lower Hadar For-
mation. Galili proboscideans are primarily grazers, but
browsing rhino (Diceros) and giraffe also are present, and
bovids are dominated by tragelaphins followed by bovins
52 A. K. Behrensmeyer and K. E. Reed
and reduncins. The habitat is reconstructed as primarily
woodland to bushland, although open grassland is indi-
cated by the grazing proboscideans and equids (Kullmer
et al. 2008).
? Fejej, Ethiopia. Although originally described as A.
afarensis, Van Couvering (2000) suggests that these
specimens may be A. anamensis, but only based on their
age (Kappelman et al. 1996).
A. anamensis summary. Faunal and other paleoecologi-
cal evidence from seven different areas indicate a range of
habitats from closed woodland (Assa Issie) to open grass-
land (Kanapoi). Wynn?s (2000) assessment that this homi-
nin ??thrived in varied ecosystems?? seems appropriate based
on current evidence. As discussed in Haile-Selassie et al.
(2010a), the mammalian species recovered in the Woranso-
Mille are different from those at Kanapoi, Allia Bay, and
other deposits of the approximately the same age. Whether
this is due to differences in environment or reflects a larger-
scale biogeographic phenomenon requires further study.
A. afarensis
? Lothagam, Kenya. Four isolated teeth found in the flu-
vially deposited Kaiyumung Member of the Nachakui
Formation, dated at *3.5 Ma, have been attributed by
Leakey and Walker (2003) to Australopithecus cf. A.
afarensis. The dominant bovid tribes of this member,
aepycerotins, alcelaphins, and bovins, indicate relatively
open and seasonally dry conditions (Harris et al. 2003).
This interpretation is supported by a decrease in Colo-
binae and an increase in Theropithecus relative to the
underlying Apak Member.
? Laetoli, Tanzania. Australopithecus fossils are relatively
rare in the Laetolil deposits in Tanzania. According to Su
and Harrison (2008), the Laetoli environment during
Austrolopithecus? times was a mosaic of woodland,
shrubland, and grassland with ephemeral streams and/or
ponds. In contrast, Kovarovic and Andrews (2007)
reconstruct it towards the wooded end of the savanna
spectrum, i.e., a mosaic of dense woodland and bushland.
In either case, there are no aquatic animals, and thus no
evidence of permanent water, which may have contrib-
uted to low numbers of A. afarensis on the landscape as
well as in the fossil assemblages.
? Woranso-Mille, Ethiopia. Haile-Selassie et al. (2010a)
describe a partial skeleton of A. afarensis from the Korsi
Dora vertebrate locality that has an estimated age of
*3.58 Ma. Additional fragmentary hominin remains are
assigned to A. afarensis but also bear traits of A. anam-
ensis. Over 1500 vertebrate specimens from this
paleontological study area (Haile-Selassie et al. 2010b)
indicate a mix of riverine forest, open woodland and
grassland habitats (see discussion under A. anamensis).
? Dikika, Ethiopia. This locality has sediments of the Basal
and lower Sidi Hakoma members of the Hadar Forma-
tion. Wynn et al. (2006) suggest that the fossils of A.
afarensis are associated with a delta and a wooded
environment, although certain species indicating open
grasslands were also present. This site may have cut
marked bones, which are controversial but if confirmed
would show that this species incorporated meat or animal
products into its diet (McPherron et al. 2010; for alter-
native viewpoint see Dominguez-Rodrigo et al. 2010).
? Hadar, Ethiopia. A. afarensis occurs in three successive
members of the Hadar Formation, persisting through
*500 kyr in spite of shifts in the fauna and vegetation
(Bonnefille et al. 2004; Campisano 2007).
? Sidi Hakoma Member. The Sidi Hakoma deposits
range in time from *3.42?3.26 Ma (Campisano 2007).
The deposits in the lowermost part of the unit indicate
higher annual rainfall and less seasonal environments
than found in any other Hadar sub-member (Reed
2008). The rest of the Sidi Hakoma Member fluctuates
between bushland and open woodland with a riverine
component until the top of the member when there is a
transgression of paleolake Hadar into the collection
areas.
? Denen Dora Member. The entire Denen Dora Member
encompasses only about 56 kyr (Campisano 2007).
There is a major increase in the abundance of redun-
cine bovids in the middle part of this time period,
indicating extensive wetland and floodplain habitat.
After this episode, there is faunal evidence for open
wooded grassland (Campisano et al. 2004; Behrens-
meyer 2008; Reed 2008) (see earlier discussion of the
A.L. 333 locality).
? Kada Hadar Member. There are two collection units
that encompass *3.2?2.94 Ma separated by the Bou-
roukie Tuff 1 (BKT-1) at *3.12 Ma (Campisano
2007). The separation is important as the habitats shift
from open woodland with some edaphic grassland to
more arid and scrub woodland habitats. The KH-2
fauna also has high proportions of antilopin and alc-
elaphin bovids, which indicate more arid environments
(Vrba 1975), especially when contrasted with other
Hadar Formation sub-members (Reed 2008).
? Ledi-Geraru, Ethiopia. Two A. afarensis molars were
recovered from the Denen Dora Member of the Hadar
Formation (Wood 2011). They were recovered with re-
duncin bovids indicating a lakeshore environment, as
well as antilopins and alcelaphins that indicate more
shrubland and grassland habitats (Reed et al., in
preparation).
4 The Habitats of Australopithecus 53
? Maka and Belohdelie, Ethiopia. White et al. (1993)
conclude from faunal evidence that there was woodland-
bushland at the time of deposition of the Maka material,
which is similar to the faunal interpretation for the Denen
Dora Member of the Hadar Formation. A. afarensis has
also been assigned to the frontal from Belohdelie, but no
information is available for the ecological context of the
find (Asfaw 1987).
? East Turkana (Koobi Fora), Kenya. The older deposits
(Tulu Bor and Lokochot members) have a moderately
large faunal collection but Australopithecus is rare. The
Tulu Bor Member of the Koobi Fora Formation is con-
temporaneous with the entire Hadar Formation in time
(3.4?2.7 Ma) but has yielded only a few A. afarensis
specimens (Kimbel 1988; Campisano et al. 2004). Feibel
et al. (1991) described the depositional environment
during Tulu Bor times as fluvial with floodplain lakes.
Harris (1991) suggested that the habitat at this time
included gallery forests amid floodplains, wet grasslands
and woodlands. Controlled paleontological surveys of the
Lokochot and Tulu Bor members at East Turkana support
the comparative scarcity of Australopithecus fossils at
East Turkana (Behrensmeyer et al. 2004), suggesting that
the pattern is ecological or paleobiogeographic rather
than taphonomic.
? West Turkana, Kenya. As mentioned previously, at least
42 catalogued but unpublished specimens are assigned to
Australopithecus, and at least some of these are assigned
to A. afarensis. These derive from above the Tulu Bor
Tuff and other fauna indicates gallery forest and wood-
land (Leakey et al. 2001).
? Fejej, Ethiopia. Hominin specimens from this site were
the oldest assigned to A. afarensis at 4.0?4.2 Ma
(Kappelman et al. 1996), although some are now regar-
ded as A. anamensis (Delson et al. 2000), but this is based
solely on the age of the remains. There is no available
information on the associated fauna or paleoenvironment.
A. afarensis summary. White et al. (1993) suggested
broad habitat tolerance for A. afarensis, and the geological
and faunal evidence from *12 different localities from
northern Ethiopia to Tanzania supports this earlier assess-
ment; the fossil remains of this species are associated with
habitats ranging from relatively open grassland to wood-
land, shrubland and riparian forest. There is no evidence
that A. afarensis preferred any particular habitat, although
low relative abundance at Laetoli and scarcity at East
Turkana suggests some limits on its ecological flexibility.
Given that this species was widespread and ecologically
eurytopic (Reed 2008), then what caused its disappearance
or extinction at *2.7 Ma? This question could possibly
further examined if: (1) the parameters of the reconstructed
habitats could be refined in terms of abiotic factors (e.g.,
seasonal extremes in temperature and moisture), (2) patterns
indicating competition or niche-partitioning could be
reconstructed for other eurytopic mammalian species
coexisting with A. afarensis (e.g., via stable isotope analy-
sis) and (3) morphological changes within the lineage
(Lockwood et al. 2000) could be associated with responses
to habitat change.
Australopithecus or Paranthropus aethiopicus
? Omo (Shungura Formation), Ethiopia. Suwa et al. (1996)
assign 19 isolated hominin teeth from a total sample of 48
to this species between 3.0 and 2.0 Ma. These occur from
members C?F, i.e., between 2.9 and 2.3 Ma; later rela-
tively robust teeth are assigned to Australopithecus
(Paranthropus) boisei. This species co-occurs with a
??non-robust?? hominin, represented by teeth that could
belong to A. afarensis, A. africanus, or early Homo. The
environment was predominantly riparian forest and
woodland based on associated faunas, which lack a strong
open grassland-adapted component until after 2.0 Ma
(Bobe and Eck 2001; Bobe et al. 2002; Alemseged et al.
2003; see earlier section).
A. africanus
? Makapansgat, Member 3, South Africa. This deposit
contains an extremely large number of mammalian
specimens (greater than 30,000), of which 24 are A. af-
ricanus. The deposit was accumulated in the cave by
fossil hyaenid and porcupine species (Maguire et al.
1980). Mammalian community structure suggests that
this region was a habitat mosaic that contained riparian
woodland, bushland, and edaphic grassland (Reed 1998).
Other habitat reconstructions range from woodland (Vrba
1980) to forest (Cadman and Rayner 1989).
? Makapansgat, Member 4, South Africa. A. africanus is
represented by only three out of a total of 257 mamma-
lian specimens. Cercopithecine monkeys make up 80 %
of the collection; and the likely accumulators were birds
of prey and leopards (Reed 1996). Member 4 fossil
deposits suggests a more wooded habitat than Member 3,
but this could be a function of sample size and predation
bias rather than an actual change of habitat at the site. As
Members 3 and 4 are roughly contemporaneous, both
assemblages probably represent a similar woodland?
bushland habitat mosaic.
? Sterkfontein, Member 2, South Africa. The skeleton of
Stw 573 has been attributed, thus far, to Australopithecus
sp. but is still embedded in rock, preventing thorough
54 A. K. Behrensmeyer and K. E. Reed
taxonomic analysis (Clarke 1999). Dating for the locality
ranges from 2.8 to 2.6 Ma (Pickering and Kramers 2010).
The fauna recovered with Stw 573 thus far is mostly
cercopithecoids and carnivores with very few ungulates
(Pickering et al. 2004). These researchers suggest an open
woodland habitat in a valley setting surrounded by rolling
hills covered with rocks and shrubs. A riverine forest is
also proposed based on the presence of numerous mon-
keys and a leopard. The fauna, other than the hyaenid
Chasmaporthetes, is also present at Sterkfontein Member
4 and other younger localities in South Africa. Thus, if
the deposit overlaps in time with Sterkfontein Member 4
(see below), there may be two Australopithecus species
present at roughly the same time. It is worth noting that
Pickering et al. (2004) state that most of the fauna
recovered are ??climbers?? and this may have implications
for Stw 573 as well.
? Sterkfontein, Member 4, South Africa. This member has
been dated to between 2.2 and 2.6 Ma (Herries et al.
2013). The faunal community suggests a habitat of open
woodland, with bushland and thicket areas (Reed 1997).
Other habitat reconstructions of this member at Sterk-
fontein have indicated medium density woodland (Vrba
1975) and an ecotone between dry sandy highveld
grassland and Kalahari thornveld (Avery 2001). Bamford
(1999) notes the presence of lianas, which indicate fairly
dense riverine forest.
? Taung and Gladysvale, South Africa. The single speci-
men of A. africanus from Taung was likely incorporated
into a meal of a bird of prey. The eagles suggested as the
predator range in their hunting regions from forests
through deserts (Berger and Clarke 1995). The other
fauna associated with this deposit suggests a habitat that
is fairly dense woodland (e.g., Tragelaphus, Cephalo-
phus, Panthera, cercopithecoids). The hominin teeth
recovered from Gladysvale are associated with other
fauna recovered from the ex situ material that indicate
deposition during a period of relatively wet climate and
closed vegetation (Berger and Tobias 1994; Plug and
Keyser 1994).
A. africanus summary. Although there is some evidence
for closed forest habitats (e.g., fossil wood, lianas), the
associated fauna recovered with this species indicates a
mosaic of habitats ranging from forest to open grassland.
Certainly the higher latitude of these deposits means
important climatic differences compared with those nearer
to the equator, especially with respect to seasonal temper-
ature fluctuations. Also, the irregular upland terrain of the
South African limestone plateau contrasts with the lower,
more even topography of the aggrading rift basins in East
Africa
A. sediba
This species, recently discovered at the site of Malapa in
South Africa, is represented by relatively complete remains
of a number of juvenile and adult specimens from a cave fill
dated to *1.9 Ma. Thus far, no other fauna has been
published from the locality, but remains of other species are
present, and information on the paleoecology will no doubt
be forth-coming (Berger et al. 2010; de Ruiter et al. 2013;
Dirks et al. 2010).
Kenyanthropus platyops
This taxon was recovered from the Nachukui Formation on
the west side of Lake Turkana (Leakey et al. 2001).
According to these researchers, fauna recovered near the
specimens suggest a habitat that is more wet and closed than
habitats at Hadar. We include this taxon because, though
not placed in the genus Australopithecus, it is from the same
time interval as early Australopithecus in East Africa.
A. garhi
? Bouri, Ethiopia. This species has been recovered from the
Hata Member of the Bouri Formation, and is a late East
African (2.5?2.1? Ma) representative of the genus (As-
faw et al. 1999). The fauna associated with A. garhi
indicates the presence of a shallow lake surrounded by
grasslands (de Heinzelin et al. 1999). Cut marked bones
were found in the same strata as A. garhi, and meat-eating
behavior is attributed to this species (de Heinzelin et al.
1999). White (2002) has suggested that some of the teeth
from the Omo Shungura Formation are A. garhi and as
such would be found in the more closed woodland hab-
itats of the region (Bobe and Eck 2001; Bobe et al. 2002;
Alemseged et al. 2003).
Discussion: Australopithecus Paleoecology
Returning to the questions that were posed at the beginning of
this paper, what can we say about the paleoecology of aus-
tralopiths in light of current taphonomic, paleontological, and
geological information from the many known occurrences of
Australopithecus in the African fossil record?
4 The Habitats of Australopithecus 55
1. What was the range of habitats associated with Austra-
lopithecus, and is it possible to discern each species?
preferred habitat? Even the earliest records for the genus
include evidence for diverse habitats, from forests and
woodlands to more open vegetation, suggesting eury-
topic ecological adaptations from the beginning. This
evidence is time-averaged over ecologically long time
intervals, thus limiting what we can infer about habitat
preferences within the available vegetation mosaics. We
also do not yet know whether any of the species in this
genus preferred one of these habitat types or a mix of
open and closed habitats. However, Campisano (2007)
has shown that across similar time intervals at Hadar, A.
afarensis is more abundant in drier regions. The docu-
mented existence of Australopithecus from Chad to
Ethiopia to South Africa indicates continent-scale dis-
tribution, considerable seasonal temperature tolerance,
and adaptability to different topographic settings.
2. What were the most important limiting ecological vari-
ables (e.g., food, water, shelter, competition with other
species, predator avoidance, intra-species interactions)
for the australopiths? There is a possibility that the
genus was limited by climatic conditions and associated
vegetation types that disappeared at Hadar during the 2.8
?2.35 Ma interval of increased aridity, when it went
locally extinct. Relative scarcity of A. afarensis fossils at
Laetoli suggests dependence on water sources and veg-
etation associated with water. Otherwise, understanding
of these variables remains unknown.
3. How did habitats vary among australopith species and
across different regions? There is evidence from regional
faunal differences for some degree of either habitat
variability or biogeographic isolation among the differ-
ent species. Contrary to the hypothesis that hominin
evolution is linked with retreating forests and expanding
grasslands, the habitats of the earliest species, A.
anamensis have been reconstructed as rather open, fol-
lowed by a mosaic of open and closed habitats for A.
afarensis. A. bahrelghazali appears to have existed in the
most open grassland habitat, which is interesting con-
sidering its location in a lake basin in central Africa. A.
africanus appears also to have been associated with
mosaic habitats, although the habitats contributing to the
mosaic change through time in southern Africa. There is
as yet no overlap in species between South and East
Africa during the temporal range of Australopithecus,
evidence that this hominin genus was one of the most
widely distributed members of the Pliocene mammalian
fauna of Africa. Differences in its patterns of occurrence
among basins within East Africa and, indeed, among
localities on the west and east side of Lake Turkana, also
suggest that Australopithecus was a eurytopic genus.
These observations and supporting data provide a
foundation for developing and testing hypotheses
regarding responses to climate change experienced on
local and regional scales. New research to obtain high
resolution drill core records of environmental change
from Plio-Pleistocene paleolakes along the East African
Rift can also be applied to these hypotheses.
4. Was there more than one Australopithecus species
sharing an ecosystem at any given place and time? This
appears possible given the evidence from West Turkana,
Omo, Galili, Woranso-Mille, and Sterkfontein
(Table 4.1), but at present there is hard evidence for only
one species at any one stratigraphic level and site. Time-
averaging of hominin remains from different time peri-
ods may create the appearance of co-occurrence in a
paleocommunity. Further fieldwork and taxonomic
research are needed on deposits that may include dif-
ferent hominins.
5. Was Australopithecus a maker and user of stone tools?
There are tantalizing occurrences of purported cutmarks
on bones at two Australopithecus sites, Dikika and Bo-
uri, but these finds are contested. More in situ evidence is
needed, including the artifacts themselves, to provide a
definitive answer to this question.
6. Was there change in habitat use through time? Did niche
breadth increase or decrease within individual lineages
as Australopithecus evolved? We do not know the
answers yet, but higher resolution paleoecological
research, additional hominin sites, and stable isotope
studies of hominin enamel through sequences such as the
Hadar Formation could provide new information bearing
on these questions.
7. How might global or continental-scale climate change
between 4.5 and 2.0 Ma have affected the paleoecology
of Australopithecus? Some degree of climate forcing is
probable, but understanding this will take careful study
of regional variability in paleoclimates in southern ver-
sus eastern versus central Africa and comparisons with
deep sea and continental lake records of global and
continental-scale climate change. These data, in turn, can
be used in paleoclimatic models of more localized
regions to arrive at better models of climatic change
through the Pliocene.
Habitats
The localities where the different species of Australopithe-
cus have been documented provide evidence for varying
amounts of closed woodland to forest as well as open
grassland and shrubland habitats. This evidence is based
primarily on associated fauna and stable isotopes, with
56 A. K. Behrensmeyer and K. E. Reed
some input from the paleobotanical record. The genus
Australopithecus can be characterized as eurytopic because
its species are found in deposits that have faunal and iso-
topic evidence for a wide range of habitats. It is not clear,
however, whether individual species were eurytopic or
stenotopic with respect to the inferred spectrum of vegeta-
tion types because hominin sample sizes are generally too
small to show statistically significant associations with
particular ecological indicator taxa (e.g., Bobe et al. 2002).
The one exception where there is enough hominin fossil
material at one locality to begin to examine this questions is
A. afarensis at Hadar, which persists for *500 kyr though
changing environmental conditions, indicating eurytopy
with respect to these conditions (Bonnefille et al. 2004;
Reed 2008). That the microwear of A. afarensis indicates
little variability in diet (Grine et al. 2006a, b), however,
may indicate that although the species inhabited different
environments, it ate something similar in all of them (see
below).
Given limited samples of hominins and known biases
introduced by taphonomic processes, pinning down an
association of a particular hominin species with a ??pre-
ferred?? habitat may be possible using quantitative analysis
associations with ecological indicator taxa. Progress in this
approach will require more data points consisting of care-
fully controlled associations of hominins and faunal or other
proxies to allow higher temporal and spatial resolution of
the consistency of these associations. Growth in under-
standing ecological indicator species associated, or not
associated, with hominins will also help this approach.
Autecological investigations including expanded stable
isotope analysis of hominin tooth enamel using minimally
destructive laser-ablation technology could also provide
direct evidence of dietary preferences and variability. Mi-
crowear and anatomical traits indicating adaptation for
climbing, walking, etc., could also support higher resolution
inferences about preferred habitats.
Diet and Food Procurement
There has been recent research that sheds light on the diet of
some Australopithcus species but also brings up further
questions. It has long been known that A. africanus mi-
crowear indicates a variable diet, but not as variable as
Paranthropus robustus recovered from the same geographic
region. In contrast, A. anamensis and A. afarensis appear to
have been more limited in their selection of foods due to the
low variation in the fine scratches that appear on their teeth
through time (Grine et al. 2006a, b). Stable isotopes of
Australopithcus taxa are discussed in Sponheimer (2013)
and Grine et al. (2012), but indicate both C3 and C4 plants
were consumed. Finally, evidence suggests that some of
these hominins may have been consuming meat or marrow
(de Heinzelin et al. 1999; McPherron et al. 2010). Thus,
while there is interesting autecological evidence provided
for many of these taxa, there are still many questions as to
how they were utilizing their habitats.
Conclusion and Future Research
We know much more about the paleobiology of Austra-
lopithecus than we did 30 years ago, and in spite of taph-
onomic and time-averaging caveats, the large number of
documented sites now provides convincing evidence that
the genus had an impressive breadth of tolerance for varied
habitats and climates. Better-coordinated research in faunal
analysis, habitat reconstruction, spatial distribution, and
taphonomic biases of the hominin fossil record at local,
regional, continental, and global scales, as well as addi-
tional new sites, should greatly expand this knowledge in
the coming decades.
Used in conjunction with species turnover patterns and
evidence for abiotic environmental change, the evidence
provided in this paper can serve as a baseline for continuing
research on the ecological context of hominin evolution.
Further advances in habitat reconstruction for Australopi-
thecus will depend on careful attention to the scale of the
evidence (temporal and spatial) versus the scale of the
desired reconstruction. Particular caution is needed to avoid
interpreting ecological features of a time-averaged faunal
list as a ??snapshot?? (single time-plane) sample of the
habitat of Australopithecus or any other hominin.
In a succession of fossiliferous strata, we usually are
dealing with varying proportions of different habitats (e.g.,
closed vs. open, or wetter vs. drier habitats) rather than the
extremes of one or the other. How these habitat ??mosaics??
are recorded in the fossil record depends on the spatial scale
of the sample as well as the amount of time represented.
Shifts of an ecotone across a depositional area through time
can also result in a similar mixed habitat signal (Behrens-
meyer et al. 2007). There is no simple solution to the
problem of time-averaged ecological signals, but in some
fossil-bearing sequences there are ways to calibrate the
scale of habitat patches and evaluate the adaptations of
individual species. These include:
1. Higher resolution sampling and morphological analysis
of faunas associated with Australopithecus-bearing
strata, including intra- and inter-basin comparisons of
mammalian species associated most commonly with
Australopithecus. What are the morphological and
abundance similarities and differences among species
that co-occur, or do not co-occur, with Australopithecus?
4 The Habitats of Australopithecus 57
To the same end, analyses of tooth wear patterns and
stable isotopes in the same species across space and/or
through time at individual localities will give us infor-
mation regarding diets that may be consistently different
in particular basins.
2. Coordinated lateral sampling of faunas and paleoenvi-
ronmental variables in Pliocene sequences where Aus-
tralopithecus is common versus uncommon or absent
(e.g., Hadar vs. Turkana Basin, Omo Shungura vs. Tugen
Hills vs. Lothagam).
While pursuing increased resolution and refinement of
taphonomic and ecological evidence, it also will be
important to adjust the spatial and/or temporal scale of
paleoecological interpretations to take account of the
inevitable limitations of the record. Much remains to be
learned about resolving ecological information in the fossil
record of Australopithecus, or any other intriguing extinct
mammalian genus.
Acknowledgments The authors thank John Fleagle for his encour-
agement and patience as well as his insightful comments on the
manuscript. We are very grateful to Richard Leakey and Lawrence
Martin for providing the focus on Australopithecus at the 2007 con-
ference at Stony Brook and for organizing a very informative and
enjoyable meeting. This is Evolution of Terrestrial Ecosystems pub-
lication #289.
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