ArticlePDF Available

Pliocene Animal Trackways at Laetoli: Research and Conservation Potential


Abstract and Figures

Laetoli, a paleoanthropological site in Northern Tanzania, is perhaps best known for its famous fossil hominid footprints that were discovered by Mary Leakey and her co-workers in 1978. The site not only preserves the hominid footprints but also trackways, which provide a snapshot of Pliocene faunal communities from East Africa and their inferred environments. Unlike the hominid footprints at site G, which have received tremendous attention, the animal trackways, especially at Localities 7, 8 and 10 have been neglected and are fast disappearing. In this paper, we discuss animal tracks at a newly discovered exposure and provide preliminary data on the tracks at this exposure and other sites. We also discuss the importance of the animal trackways as ecological indicators, which we have investigated as part of ongoing research and conservation efforts initiated by the Tanzania Field School in Paleoanthropology and the Associated Colleges of the Midwest (ACM) Tanzania Semester Abroad programs.
Content may be subject to copyright.
This article was downloaded by: [Auraria Library]
On: 10 August 2014, At: 15:27
Publisher: Taylor & Francis
Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House,
37-41 Mortimer Street, London W1T 3JH, UK
Ichnos: An International Journal for Plant and Animal
Publication details, including instructions for authors and subscription information:
Pliocene Animal Trackways at Laetoli: Research and
Conservation Potential
Charles M. Musiba
, Audax Mabula
, Marie Selvaggio
& Cassian C. Magori
Department of Anthropology , University of Colorado , Denver, Colorado, USA
Department of History , University of Dar Es Salaam , Dar Es Salaam, Tanzania
Department of Anthropology , Southern Connecticut State University , New Haven,
Connecticut, USA
Department of Anatomy and Histology , Bugando University College of Health Sciences ,
Mwanza, Tanzania
Published online: 02 Dec 2008.
To cite this article: Charles M. Musiba , Audax Mabula , Marie Selvaggio & Cassian C. Magori (2008) Pliocene Animal Trackways
at Laetoli: Research and Conservation Potential, Ichnos: An International Journal for Plant and Animal Traces, 15:3-4,
166-178, DOI: 10.1080/10420940802470383
To link to this article:
Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained
in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no
representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the
Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and
are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and
should be independently verified with primary sources of information. Taylor and Francis shall not be liable for
any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever
or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of
the Content.
This article may be used for research, teaching, and private study purposes. Any substantial or systematic
reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any
form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http://
Ichnos, 15:166–178, 2008
Taylor & Francis Group, LLC
ISSN: 1042-0940 print / 1563-5236 online
DOI: 10.1080/10420940802470383
Pliocene Animal Trackways at Laetoli: Research and
Conservation Potential
Charles M. Musiba,
Audax Mabula,
Marie Selvaggio,
and Cassian C. Magori
Department of Anthropology, University of Colorado, Denver, Colorado, USA
Department of History, University of Dar Es Salaam, Dar Es Salaam, Tanzania
Department of Anthropology, Southern Connecticut State University, New Haven, Connecticut, USA
Department of Anatomy and Histology, Bugando University College of Health Sciences,
Mwanza, Tanzania
Laetoli, a paleoanthropological site in Northern Tanzania, is
perhaps best known for its famous fossil hominid footprints that
were discovered by Mary Leakey and her co-workers in 1978.
The site not only preserves the hominid footprints but also track-
ways, which provide a snapshot of Pliocene faunal communities
from East Africa and their inferred environments. Unlike the
hominid footprints at site G, which have received tremendous
attention, the animal trackways, especially at Localities 7, 8 and
10 have been neglected and are fast disappearing. In this paper,
we discuss animal tracks at a newly discovered exposure and
provide preliminary data on the tracks at this exposure and other
sites. We also discuss the importance of the animal trackways as
ecological indicators, which we have investigated as part of ongoing
research and conservation efforts initiated by the Tanzania Field
School in Paleoanthropology and the Associated Colleges of the
Midwest (ACM) Tanzania Semester Abroad programs.
Keywords Laetoli, animal trackways, paleoecology, animal track-
way conservation
The uniqueness of Laetoli as an ecologically significant
Pliocene upland site in eastern Africa makes it particularly
well suited for multifaceted paleoenvironmental studies. For
example, the Laetoli paleobiota (Table 1) and the ichnofossil
record (Table 2) contains a wide diversity of species whose
ancestors today occupy a variety of habitats ranging from open
grassland to brush and woodland (Kent, 1941; Leakey et al.,
1976; Leakey and Hay, 1979; Leakey, 1981; Hay, 1978, 1980,
1987; Ndessokia, 1990; Manega, 1993; Kyauka, 1994; Musiba,
1999). As a Pliocene site, Laetoli differs significantly from other
Address correspondence to Charles M. Musiba, Department of
Anthropology, University of Colorado at Denver and Health Sciences
Center, Campus Box 103, P.O. Box 173364, Denver, CO 80217-3364.
African paleoanthropological sites such as Hadar or Koobi Fora,
which primarily formed in fluviatile-lacustrine or floodplain
deposits (Butzer, 1974; Coppens et al., 1976; Coffing et al.,
1994; Feibel et al., 1989; Johanson and Edey, 1981; Vrba, 1975,
1985a & b, 1995, 2000; WoldeGabriel et al., 1994; Wood and
Collard, 1999). Laetoli formed entirely on Aeolian volcanic
tuffs with no major lakes or rivers. The Laetoli vertebrate
fossil fauna is also of great interest because its taxonomic
composition differs from that of other East African fauna of
comparable age (Gentry, 1981; Harris, 1985; Verdcourt, 1987).
Most remarkable are the Footprint Tuffs at Laetoli in which
tracks and trails of a great variety of animals, many of which are
long extinct and are well-preserved (Day and Wickens, 1980;
Leakey, 1981, 1987). In this paper we present some preliminary
inferences concerning Laetoli’s past ecological setting through
analyses of Laetoli’s animal trackways—their identification,
distribution, composition and significance as paleoecological
indicators—with an emphasis on the site conservation efforts
currently undertaken by our team.
The Pliocene site of Laetoli is located about 36 kilometers
south of Olduvai Gorge in northern Tanzania (Fig. 1). This
site, which covers an area about 100 km
may be viewed as
contiguous with Olduvai Side Gorge, where fossiliferous strata
of volcanic origin spanning from 4.32±0.06 to 0.21 ± 0.02 Ma
occur (Hay, 1978; Drake and Curtis, 1987; Manega, 1993).
The geology, geochronology, and stratigraphy of Laetoli with
its fossil bearing deposits and footprint horizons have been
described in detail by Hay (1978), Drake and Curtis (1978), and
Manega (1993). The footprint horizons, which are referred to as
the Footprint Tuffs, consist of ash particles mainly cemented by
calcite and less commonly by phillipsite (Hay, 1987).
Previous paleoanthropological research at Laetoli by Leakey
and her co-workers produced more than 20 fragments of
cranial and post-cranial remains of fossil hominids ranging from
Australopithecus afarensis to anatomically modern humans
(White 1977, 1980; Day and Magori, 1980). Other significant
Downloaded by [Auraria Library] at 15:27 10 August 2014
Partial list of common mammals
(nonhuman) in the fossil
record from Laetoli (Laetolil and Upper Ndolanya beds (after
Leakey, 1987; Harris, 1987; Hooijer, 1987; Gentry, 1987;
Davies, 1987).
No. of
species/genera NISP (%)
Cercopithecidae 5 91 (0.8)
Elephantidae 2 246 (2.1)
Equidae 4 371 (3.2)
Leporidae 1 992 (25.6)
Pedetidae 2 94 (1.6)
Rodentia 22 402 (3.4)
Carnivora (all taxa) 24 266 (2.3)
Rhinocerotidae 2 671 (5.6)
Giraffidae 4 897 (7.6)
Suidae 3 334 (2.7)
Bovidae 22 5248 (44.9)
Total 91 11706 (96.8)
NISP = Number of identified specimen per taxon.
The taxa listed are represented by more than 100 identifiable
discoveries at Laetoli include a trail of footprints generally
attributed to Australopithecus afarensis made by two or three
individuals and dated to 3.56 million years ago. These are
well preserved in volcanic ash within the Upper and Lower
Laetolil Beds (Leakey, 1981, 1987). Laetolil Beds is the official
term used to define the oldest deposits: the orthography is
different from the site name: Laetoli (Hay, 1987; Leakey
1987). The Footprints Tuffs, which are fine-grained volcanic ash
layers, document the world’s only undisputed footprint evidence
for the origin of habitual bipedal locomotion in the human
lineage during the Pliocene (Leakey, 1987; Tuttle, 1987, 1992).
Additionally, animal and avian tracks including raindrop prints
are also well preserved within the Footprint Tuffs. Laetoli’s
stratigraphic sequence, particularly the Ndolanya Beds also
contain numerous stone artifacts and an early manifestation of
the Middle Stone Age (MSA) or “Sangoan” traditions (Mabulla,
in preparation; Ndessokia, 1990).
Tracks and prints of animals and birds, which have been
discovered and identified at Laetoli, are known from 18 exposed
footprint horizons referred to as the Footprint Tuffs within
Localities 2, 7, 7 East, 9, 10 East, 11, 13, and 14 (Fig. 1). The
sites at these localities are listed from A–P (see Leakey, 1987 for
details). At Laetoli, the animal, avian, and insect tracks, raindrop
imprints and the fossil faunal remains are unique in terms
of their exceptional preservation. The stratigraphic sequences
and geochronology of the Laetoli site were described in detail
by Hay (1978, 1980, 1987), Drake and Curtis (1987), and
Manega (1993). The strata comprise a variety of volcanoclastic
units of varying thickness and composition. According to
Hay (1987), the source of the Laetolil Beds was a volcanic
eruption at Sadiman, which is located 15 km east of Laetoli.
The Footprint Tuffs are of eolian tuff origin (with grain size
ranging from 0.05—0.40 mm) interbedded with widespread
raindrop horizons. The tuff layers were precisely dated Drake
and Curtis (1987) and Manega (1993) using K/Ar and Ar/Ar
dating methods. Hay (1987) also produced paleomagnetic dates
complimenting the K/Ar and Ar/Ar dates, and believes that the
Footprint Tuff was deposited over a period of few weeks at the
end of the dry season and into the beginning of the rainy season.
The animal and avian trackways at Laetoli occur within the
Laetolil Beds, between the Footprint Tuffs 3 and 14 in the lower
unit and between tuffs 1 and 4 in the upper unit (Hay, 1987).
Almost immediately after the rainfall, animals, birds, insects and
hominids walked on the wet ash surface. The footprints were
buried by subsequent ash falls, allowing them to be preserved.
Later, water and wind erosion re-exposed the prints.
List of identifiable insects, birds and mammal
taxa from the footprints and trackways within the
Footprint Tuffs (after Leakey, 1987)
Insects (including termitaries)
Aves indet.
Phasianidae (guinea-fowl)
Carnivora indet.
Chalicotheridae (including Ancylotheriopus
tanzaniae ichnosp. nov.)
Simatherium kohllarseni
Hominidae gen. et sp. indet.
Bovidae (including Madoqua), and various indet.
Synonymous with Ancylotherium hennigi per
Leakey and Harris (1987).
The Laetoli hominid footprints have been attributed
to Australopithecus afarensis by many researchers.
However, the footprint structure and the inferred pedal
morphology of the Laetoli bipeds do not fit the morphol-
ogy of Australopithecus afarensis (Tuttle, 1991).
Downloaded by [Auraria Library] at 15:27 10 August 2014
FIG. 1. Map of Laetoli showing the paleoanthropological localities (numbers) and animal trackways (letters).
Unlike the hominid footprints, which have received much
attention and media publicity geared towards their conservation
in 1996–98 (Agnew and Demas, 1998; Demas et al., 1996;
Feibel et al., 1995), the animal trackways at Laetoli have been
neglected. Since Leakey and her co-worker’s detailed study of
the animal and avian tracks in 1987, no further systematic
studies of the exposed animal tracks have been conducted.
Recent observations by our teams during field school sessions
at Laetoli indicate an alarmingly rapid disappearance of the
prints that were initially documented by Leakey (1987) through
both erosion and animal trampling (Fig. 2). Unfortunately, there
seems to be no future plans for preservation and documentation
of these important ichnofossil trails. The present study was
conducted by our teams during four consecutive one month field
seasons (2002–2005), as part of ongoing summer field school
and semester abroad programs for anthropology undergraduate
students from the United States and Tanzania. The major
objectives of our study were: a) to document the existing
animal trackways as well as new exposures, b) to re-evaluate
the exposed animal and avian trackways and the footprint tuffs
in terms of their conservation needs, and c) to use the animal
trackways to construct species abundance estimates and use
that data in paleoecological interpretations of Pliocene Laetoli
(particularly within the Upper Laetolil Beds).
Recording the Animal Trackways
A total of six footprint sites were systematically surveyed,
three in each field season. Five of the surveyed sites were
previously described by Leakey et al. in 1987. These include;
site C at Locality 7, site D at Locality 11, site J at Locality
10 East, and sites L1 and L2 at Locality 13. The sixth site is
new and was brought to our attention by Mr. Simon Mataro,
a Maasai elder and guard of the hominid footprints at site G
(Locality 8). This newly discovered site was named site Q
within Locality 8, following Leakey’s (1987) site naming system
(Fig. 1).
Initial fieldwork at the animal trackway sites included setting
up transects with grid systems of 1 m × 1 m squares on a
10 m long segment of exposed footprint tuffs (Fig. 4). Then
the sites were cleared of any overburden to expose the animal
trackways for the detailed studies. The prints were cleaned using
soft brushes and brooms to reveal detailed features. Dental
and softwood picks were used to remove fine-grained debris
from the footprints, particularly prints that were selected to
be photographed. Extreme care was taken to ensure that the
footprints were not damaged in the process. Exposed prints
were mapped and identified in consultation with Mr. Mataro,
following the field guide to tracks of Stuart and Stuart (1994)
and Leakey’s 1987 initial report on identified prints.
Downloaded by [Auraria Library] at 15:27 10 August 2014
FIG. 2. (A) Maasai herders driving their livestock near Site O at Locality 14. (B) Photograph showing the extent of erosion on animal footprints at Site C
Locality 7. (See Color Plate XV)
At each site, werecorded the number of identifiable prints,
orientation, length and width of each footprint using digital
calipers and tape measures. Well-preserved prints were mapped,
and photographed and molds were made for archival purposes.
These molds are reposited in the Department of Anthropology
at the University of Colorado at Denver and Health Sciences
Center in Denver and in the Department of Antiquities in Dar
es Salaam, Tanzania. Digital photographs were made and will
be used to digitally monitor the extent of weathering process on
individual prints. At each site, datum points were established
and used for measurements to determine orientation at each
location relative to the datum point (Fig. 3). Hand-held Brunton
compasses were used to measure the angles while tape measures
were used to determine distances between prints.
Every print in our study was given a label. The first part
of the label identifies the locality and the second part is a
number that identifies the order in which the print was found.
For example, the third print identified at Locality 8 was labeled
L8P3 (L8 for Locality 8, and P3 for Print # 3). Due to the
fact that many animal trackways at various sites at Laetoli are
disappearing, at site Q we decided to excavate and record as
many prints as possible. We excavated a portion of the footprint
tuff at this site, in an area that was covered with packed-down
dirt and sand). First we removed the matrix using trowels and
brushes and cleared away the overburden covering the prints.
This exercise was done for two main reasons: first to determine
the condition of the prints and second to identify individual
Downloaded by [Auraria Library] at 15:27 10 August 2014
FIG. 3. Photographs (A & B) showing the established grid system during the documentation and mapping of the animal prints at Site D at Locality 11. (See
Color Plate XVI)
Reconstructing the Laetoli Paleoecology
from the Track Record
The partialreconstruction of the Pliocene Laetoli landscape
using ichnofossil data focused mainly on a comparative
approach that used habitat equivalence and behavioral ecology
of extant animals of the Serengeti Plains. Assuming that Laetoli
animals utilized the landscape in a way similar to their extant
counterparts, our goal was to reconstruct the paleoguilds and
establish possible habitat preferences of the print makers.
Using Kingdon’s (1997) Guide to Mammals of East Africa,
we constructed Laetoli’s Pliocene paleoguilds and established
habitat preferences for each of the animals represented in the
trackways. The data we collected do not necessarily provide
a complete picture of Laetoli’s paleolandscape; however, they
provide us with snapshots of the landscapes that the print-
makers experienced 3.8 million years ago. Therefore, our
paleoenvironmental analysis is currently restricted to qualitative
interpretation of Laetoli’s ancient ecology.
Our preliminary results on the reconstruction of Laetoli mam-
malian and avian paleoguilds is based on qualitative data and
thus restricted in its interpretation to general ecological trends.
The results presented below also provide brief descriptions of
the individual sites that we studied.
Sites Q at Locality 8
Site Q which was discovered in 2003, consists of a small
exposure of only a few square meters. Preservation conditions
of exposed prints on this site are relatively poor and most of
the prints are extremely eroded. We were able to tentatively
identify 28 individual prints that were exposed. The prints are
of ancient species of giraffe, roan antelope, gazelle, guinea fowl,
rhino and buffalo (Table 3A). Conspicuously absent at this site
are Madoqua and lagomorph prints, which are plentiful at other
sites we studied.
Physiographically site Q is located in a channelized gully
undercut by a seasonal stream that erodes and exposes the
footprint tuffs. The majorities of the guinea fowl prints at this
site are extremely eroded and may not survive the torrential
rains of the next few years. Unfortunately, if no action is taken
to preserve these prints it is clear that they will soon disappear.
Furthermore, roots of thorny acacias trees have penetrated and
shattered some of the trackways, while daily herding of goats
List of identified animal prints/per taxon at Sites Q and R.
Identifiable species
No. of identifiable
% of total
identified prints
A. Site Q
Giraffe 5 17.86
Roan Antelope 4 14.29
Gazelle 3 10.71
Guinea Fowl 13 46.43
Rhinoceros 2 7.14
Buffalo 1 3.57
Total 28 100
B. Site R
Elephant 8 0.44
Madoqua 858 47.69
Lagomorpha 859 47.74
Buffalo 4 0.22
Giraffe 11 0.61
Carnivore indet. 9 0.56
Lion 2 0.11
Baboon 3 0.17
Rhinoceros 7 0.39
Eland 4 0.22
Gazelle 12 0.67
Hartebeest 8 0.44
Roan Antelope 7 0.39
Beetle 1 0.06
Guinea Fowl 5 0.28
Aves indet. 1 0.06
Total 1,799 100
Downloaded by [Auraria Library] at 15:27 10 August 2014
FIG. 4. Photograph showing a 3 × 3 meters excavated trench of on the Footprint Tuffs at Site C, Locality 7. (See Color Plate XVII)
and cattle through thesite exert tremendous damage to the
exposed prints (Fig. 2 and 4). At one end of the exposure, the
tuffs have completely been obliterated and the prints washed or
weathered away. Some of the tuffs at this site however, are still
covered by tightly packed and well sorted matrix.
Site R at Locality 7
Site R at Locality 7 consists of a large exposure of the
footprint tuffs that lie within a seasonal streambed. Large
portions of the exposed tuffs have been eroded, leaving behind
few recognizable prints. Some areas of the exposed tuffs still
harbor many identifiable prints. At this site we were able to
identify 1,799 prints, where a large portion of the prints were
assigned to lagomorphs and Madoqua. Other identifiable prints
represented ancient elephants, buffalos, giraffes, lions, baboons,
rhinoceros, elands, gazelles, hartebeests, roan antelopes, guinea
fowls and beetles. A number of indeterminate carnivore and bird
prints were also found at the site (Table 3B).
Current preservation conditions of the prints at this site are
similar to that of site Q; cattle and goats have frequently been
herded across the exposed tuffs thus severely damaging the
Sites D at Locality 11
Site D is the largest animal trackway site at Laetoli with
exposed footprint tuffs containing many trails representing
elephants, hyenas and guinea fowl. This site is one of the few
sites where Leakey (1987) extensively documented the animal
trackways. Since then more tuffs have been exposed, and we
were able to identify two of the three hyena trails as well as an
elephant trail that was initially described by Leakey in 1987.
At this site we identified a large number of prints comprising
a wide variety of animals. Again, Lagomorphs and Madoqua
prints dominated the footprint assemblage at Locality D (Table
4A). Additionally, we were able to identify five prints made by
Simatherium kohllarseni, a large extinct bovid.
Large portions of exposed footprint tuffs at the site are
crumbling away due to water erosion and livestock trampling
(Fig. 5). We observed a series of fractures of an evaporitic
nature that run through the footprint tuffs. It was not unusual
to see donkeys, cows or goats milling around on top of the
exposed prints. During the rainy season water puddles form in
large animal prints providing drinking water to goats and other
animals. The Maasai who live nearby also collect rainwater from
the puddles, scraping the prints with plastic buckets and other
Downloaded by [Auraria Library] at 15:27 10 August 2014
List of identified animal prints/per taxon at Sites D and J.
Identifiable species
No. of
% of total
identified prints
A. Site D
Elephants 11 0.80
Madoqua 545 43.18
Lagomorpha 544 43.10
Buffalo 7 0.60
Carnivore indet. 5 0.40
Baboons 7 0.60
Gazelle 2 0.10
Hartebeest 3 0.20
Hyena 52 4.10
Guinea fowls 70 5.50
Aves indet. 2 0.10
Totals 1262 100
B. Site J
Elephant 16 12.31
Madoqua 66 50.77
Buffalo 4 3.07
Hyena 5 3.85
Guinea Fowl 37 28.46
Zebra 2 1.54
Total 130 100
Site J at Locality 10 East
Site J lies within a seasonal streambed at Locality 10 East. As
a result, large portions of the exposed tuffs have eroded away,
leaving few recognizable prints. Other areas of the exposures
do contain many identifiable prints, however; we were able
to identify 130 prints, which were dominated by lagomorphs
and Madoqua (Table 4B). The site contains prints of several
large bovids, giraffids and hyenas. Interestingly enough, the
directionality of the giraffe prints are oriented toward the south
while those of zebra, buffalo and hyenas are oriented toward the
northwest direction. Rhino prints are remarkably absent at this
site when compared with site R at Locality 7 (Leakey, 1987).
Sites L1 and L2 at Locality 13
The exposed footprint tuffs at locality 13 measure approx-
imately 68 m
on a U-shaped outcrop with an open end to
the south (Leakey, 1987). The area around this site is flat
covered with grasses and isolated thickets. Therefore, the site
is vulnerable to Maasai cattle trampling and erosion by water.
Consequently, large parts of the footprints at this site have been
damaged. Two areas with apparent prints were named as Site L1
and L2, respectively. Site L2 occurs about 50 meters west of L1.
At both sites, rhinoceros prints predominate among those of the
large animals (Tables 5A and B). At L1 site, all rhinoceros prints
have a northwest orientation, while at L2 site some trackways
head northwest and others toward the east. Raindrop imprints
are numerous at site L1.
Attempts to construct Laetoli mammalian paleoguilds based
on footprint identification produced 11 mammalian and one
avian family. These families are: Hyaenidae, Elephantidae,
Leporidae, Bovidae, Giraffidae, Rhinocerotidae, Bovidae, Cer-
copithecidae, Felidae, Equidae and Numididae. We were able
to tentatively identify tracks made by seventeen different types
of animals and birds: giraffes, roan antelopes, gazelles, guinea
fowls, rhinoceroses, buffalos, elephants,baboons, hyenas, lions,
rabbits, dik-diks, oribi, hartebeests, elands, zebras and the now
FIG. 5. (A) Photograph showing the eroded Footprint Tuffs at Site C, Locality 7 before conservation efforts. (B) Photograph showing the boulder-column
supporting the heavily eroded Footprint Tuffs at Site C. (See Color Plate XVIII)
Downloaded by [Auraria Library] at 15:27 10 August 2014
List of identified animal prints/per taxon at Sites L1 and
Identifiable species
No. of
% of total
identified prints
A. Site L1:
Lagomorph 15 25.42
Madoqua 15 25.42
Rhinoceros 14 23.73
Lion 1 1.70
Guinea Fowl 10 17.00
Zebra 3 5.08
Indeterminate 1 1.70
Total 59 100
Site L2:
Elephant 4 13.30
Rhinoceros 18 60.00
Guinea Fowl 6 20.00
Hyena 2 6.70
Total 30 100
extinct bovid species of Simatherium kohllarseni. Overall, rabbit
and dik-dik prints were the most numerous, followed by guinea
fowl, hyena, rhinoceros, elephant, giraffe and gazelle of varying
The two main processes responsible for the footprint
destruction at Laetoli are of natural and anthropogenic nature.
Natural processes include erosion and weathering mainly caused
by rain and wind. Also a series of fractures composed of
carbonate in-fills as well as tree roots were noted at almost
all sites. Water erosion occurs mainly during the wet season and
heavily water-worked and eroded tuffs are commonly present.
Wind erosion, especially during the dry season (May throughout
August) is common at Laetoli, and its effects can easily be
recognized on exposed tuffs. Anthropogenic agents include
livestock herding and scouring for water during the rainy season.
The Maasai who live in the area practice pastoralism and drive
their cattle over the exposed tuffs on a daily basis. The animal’s
hooves over time cause considerable damages to the footprints.
Past paleoecological studies at Laetoli focused predomi-
nantly on faunal remains, particularly those of fossil ungu-
lates, fossil pollens and paleosols. The ecological information
available from the animal trackways has not been an im-
portant component of any past environmental reconstruction.
For example, Hay (1980) used fossil faunal assemblages to
postulate that Laetoli’s Pliocene environs closely resembled
open country savanna similar to modern environments in the
modern Serengeti Plains. Bonnefille and Riolett’s (1987) study
using fossil pollen also supported this view. Harris (1985), on
the other hand, proposed a closed habitat of mixed forest for
Pliocene Laetoli. Other studies using similar evidence have
come to different conclusions. Gentry (1987) suggested that
Laetoli paleolandscapes were composed of brush or thorny
shrub habitats contra to Denys (1987), who found evidence
for wooded brush/savanna grasslands. Most recently, a study
of bovid limb morphology from Laetoli fauna conducted by
Musiba (1999) and Musiba and Magori (2006) indicate that
Pliocene Laetoli was a mosaic environment composed of
grassland, light cover and galleries of woodland, much like
the modern Tarangire National Park. The ecological snapshot
that the trackways provide can enhance our understanding
of Laetoli’s past environments and allow testing of different
ecological hypotheses.
Preliminary analyses of the presence and absence of terres-
trial mammalian guilds at Laetoli (Table 6) and their preferable
habitats (Table 7) indicate that Laetoli landscape 3.56 Mya
was very mosaic and differed significantly from the present
Serengeti landscape. The presence of Simatherium Kohllarseni,
Madoqua, giraffe, and roan antelopes strongly indicate the
existence of closed habitats characterized by open woodlands,
wooded grasslands and seasonal floodplains. Furthermore, the
presence of Guinea fowls, rhinoceros, buffaloes, baboons,
hyenas, Madoqua, hartebeests and elands suggest that Laetoli
was mosaic, characterized by galleries of trees, open grasslands,
woodlands, shrubs and thickets.
As predicted, the types of animal tracks we found are similar
to those that Mary Leakey’s team uncovered (Leakey, 1987). In
fact, she found prints made by all of the same animals that we
did. However, Mary Leakey also found tracks made by animals
that we did not find, including Struthianidae, Hominidae and
Suidae as well as Hipparion and Chalicothere prints. Our limited
discoveries, in comparison to Leakey’s team, may be the result
of our short field season and our focus on conservation rather
than on exploration for new sites.
Working under the assumption that habitat preferences of
Pliocene animals at Laetoli would have been similar to the
habitat preferences of their extant counterparts, we are able to
construct general habitat preferences for each inferred animal
represented by prints in the footprint tuffs at Laetoli (Table 6).
Extant counterparts of most of the animals represented by
tracks in the footprint tuffs prefer to live in areas where trees
are present rather than in areas of open savanna grassland. For
example, the predominance of Madoqua (Dik-dik) prints is in-
dicative of a mosaic environment (Musiba, 1999). Furthermore,
Madoqua prints are common at Laetoli and were found at four
out of the six sites we studied, thus indicating that patches of
woodlands might have existed at Laetoli during the Pliocene.
The presence of bovids, especially elands and buffalos as well as
giraffes, and Simatherium kohllarseni also indicate that Pliocene
Laetoli consisted of several different micro-habitats. If the
Downloaded by [Auraria Library] at 15:27 10 August 2014
List of reconstructed Laetoli mammalian paleoguilds with
number of identifiable prints/taxon.
Identifiable species
No. of identifiable
% of total
identified prints
Elephant 39 1.18
Lagomorphs 1450 43.91
Madoqua 1449 43.88
Buffalo 12 0.36
Giraffe 16 0.50
Rhinoceros 41 1.24
Gazelle 17 0.51
Hartebeest 11 0.33
Roan Antelope 11 0.33
Hyena 59 1.80
Guinea Fowl 141 4.27
Aves indet. 3 0.09
Baboon 10 0.30
Carnivore indet. 15 0.45
Lion 3 0.09
Eland 4 0.12
Zebra 5 0.15
Indeterminate 11 0.33
TOTAL 3302 100.00
habitat preference of modern animals is a reliable indicator
of past habitat preferences, then Pliocene Laetoli consisted
of a variety of habitats including open grassland and wooded
galleries (see also Musiba, 1999).
The Tanzania Field School has conducted a pilot conservation
project at site 7 for the past several years. The goals of this
project include:
1.) the preservation and documentation of recently exposed
animal tracks,
2.) identifying the major erosional forces operating at the site
and initiating efforts to reduce them, and
3.) encouraging the involvement of local communities in site
The major erosional forces operating at this site are similar
to those at other Laetoli sites namely, seasonal rain water,
livestock, and tree roots. However, the seasonal rains present
a specific problem at this site. A parapet constructed by Mary
Leakey to divert seasonal run-off from a portion of the site is in
disrepair. During the rainy season, water appears to flow from
the parapet to lower areas of the site. This channeling of the
water into a relatively narrow area has resulted in a scoured-out
streambed exposing several tuffs with animal tracks. Based on
our observations over the past five years, the animal tracks in
this section of the site are eroding rapidly.
In conjunction with the Tanzanian Antiquities Department,
and with the consultation and permission of local authorities and
Animals and their reconstructed habitat analogies.
Animal Preferred Habitat
Giraffe Open woodlands, wooded grasslands, seasonal floodplains
Roan antelope Wooded grassland—need tree clumps for cover
Gazelle Savanna grassland
Guinea fowl Dry areas with more or less tree cover, shrub, and bush
Rhinoceros Edges of thickets and savannahs with short woody growth
Buffalo Savanna with patches of thicket, reeds, or forest
Elephant All vegetation types, changes with season and amount of vegetation available
Baboon Needs trees for survival; woodland/savanna
Hyena Dry acacia bush, open plains, but present in wooded country
Lion Savanna and ecotone, but not closed forests and dry deserts
Lagomorph Grassland or rocky grassland
Madoqua Never seen far from cover; mosaic (Musiba 1999)
S. kohllarseni Closed cover conditions, including forest galleries
Oribi Wooded grassland
Hartebeest Ecotone between wood or scrub and open grassland
Eland Savanna +woodland
Zebra Bush/grass mosaic
Downloaded by [Auraria Library] at 15:27 10 August 2014
Maasai people living in surrounding areas, several conservation
projects were initiated to reduce erosion and document a sample
of animal tracks exposed in the streambed. Tanzanian teachers
(both high school and elementary school) and local people par-
ticipated in all aspects of the conservation projects. Over the past
several years, budget and time considerations limited our efforts
to small projects that could be completed within a few weeks.
Seasonal water flow into the streambed was found to be the
major source of erosion of the exposed tuffs. To reduce water
flow into this area, loose boulders were positioned along the
parapet in the area of disrepair. A permanent solution should
include a hydraulic assessment of the immediate area and
incorporate the needs and lifestyle of local populations. While
seasonal rains scour the surface of exposed tuffs, tree roots are
undermining both unexposed and exposed tuffs.
Acacia trees grow in profusion around the streambed. The
majority of trees appear to be young with trunk diameters of less
than three centimeters. Careful removal of topsoil at the edge
of the streambed in two 3 × 3 meter areas was undertaken to
identify the extent of root damage to the underlying tuffs (Fig.
4). One clearing was made near a mature tree while the other
was in an area of scrub brush. Several roots measuring over two
centimeters in diameter were found to be embedded in the tuffs
in each area. The embedded roots caused uplift, cracking and/or
complete erosion of areas of the tuffs. Before the clearings were
re-covered with topsoil, the roots were cut and several layers of
burlap were placed over the tuffs’ surface to inhibit further root
expansion. The availability of seasonal water and the shade of
a few mature acacia trees draw both wild and domestic animals
to the site. However, as their hooves scrape the tuffs’ surface,
they damage the ancient animal tracks. The importance of the
site and its erosional problems were discussed with the elders
of several local Maasai villages. An agreement was reached
about protecting the site. Local Maasai were hired to cut young
acacias and position them as a livestock barrier around the site.
The local people then agreed to monitor the site and to make
repairs to the barrier as needed. In addition to protecting the site
from livestock, the removal of many of the young acacia trees
from the site, may reduce the extent of root damage to the tuffs.
In order to document the site, photographs and general
information about the animal tracks continues to be collected.
While numerous, the animal tracks in the streambed exhibit
various stages of erosion. Due to the great number of exposed
tracks and the time limits of this project, only a small sample of
tracks were documented by measuring their maximum length,
width and depth, as well as their orientation and their association
with other tracks. In addition, casts were made of specific
tracks using a silicone-based material that lifts easily and is
nondestructive to the matrix. Duplicates of the casts were given
to Tanzania’s Antiquities Department. The casts and other data
we collected enhance the documentation of this site which may
disappear due to erosion.
Perhaps the greatest issue in terms of conservation is the
lack of understanding of the paleontological and paleoecological
FIG. 6. Photographs showing the progressive weathering of the baboon
footprint at Site C. Locality 7. (A) Photo taken during the 2002 field season.
(B) Photo taken three years later during the 2005 field season. (See Color Plate
importance of the animal tracks. Clearly the current unchecked
state of deterioration of this rare and rich paleontological
record will result in its rapid disappearance (Fig. 6). However,
there are several courses of action that the custodians of these
treasures could implement as part of a long term conservation
solution. Natural agents are to some extent impossible to control.
Nevertheless, if certain conservation efforts can be implemented
then we may expect to see a decrease in the deterioration of these
important indicators of the ancient environment. For example,
to combat water-based erosion, water diversion structures like
ditches could be built near the exposed footprint tuffs. Such
structures could impact the Maasai living in the area, as they may
be forced to find new trails along which to drive their livestock.
Therefore, such actions would require Maasai participation in
decision making in conservation of Laetoli in general.
As our conservation project shows, the local Maasai can
play important roles in site conservation. To maintain sites, they
could weed out seedlings that have taken root on the exposed
tuffs. They could also cut down trees like those at the hominid
Downloaded by [Auraria Library] at 15:27 10 August 2014
footprint Site G at Locality 8 that have already punched through
the tuffs. This would prevent them from destroying more of
the tuff as they continue to grow. Tree stumps could be treated
chemically to avoid damage to the tuffs.
Conservation experts who worked on the hominid footprints
did cut and remove Acacia roots using mini powered tools
and surgical instruments. Similarly, the same remedy could be
implemented for the animal trackways at Laetoli. Loosened
tuffs could be strengthened by applying soil consolidants before
removing the roots (Agnew and Demas, 2000). Furthermore,
a monitoring program as part of a comprehensive long-term
conservation program could be established in collaboration
between the local Maasai, archaeologists, and the Department
of Antiquities in Dar es Salaam.
In addition, an educational program could be implemented
which could alleviate some of the anthropogenic destruction of
the footprint tuffs. For example, as we have done, the Maasai
who live in the area could be involved and educated about
the importance of the site. Working in collaboration with the
Antiquities Department, the Ngorongoro Conservation Area
Authority and local village governments, an outreach program
can be established that will include and inform the local Maasai
about the importance of Laetoli. The animal trackway sites
could potentially be used as a cultural heritage attraction that
could bring in revenues to the community. It is clear that a
major conservation effort is required to preserve the animal
trackways. However, successful conservation will require the
involvement of local people and the development of projects
which are compatible with their lifestyle.
Major mammalian radiations, including the hominini linage,
have been influenced by global climate shifts. However, the
extent of such climate shifts and the subsequent ecological
changes on the African content are still not very well understood.
Laetoli, the only upland Pliocene paleontological site in Sub-
Saharan Africa, sports perhaps some of the most important
clues that might assist us to fully understand the effect of
Pliocene climate shift and turnover pulses. Laetoli’s animal
trackways are of great importance in that they contain snapshots
of time-averaged paleoecological events that may help us
understand high-fidelity ecological changes that are usually
hard to discern from biostratigraphic evidence. Therefore, a
paleoenvironmental reconstruction using the ichnofossil record
available at Laetoli is necessary in understanding trends
associated with human evolution during the Pliocene in East
Africa. This study, which we consider as preliminary in nature,
enhances our understanding of Laetoli’s ancient landscape
through the use of mammalian paleoguilds presented in the
ichnofossil record.
Our preliminary results, based on the animal tracks we
investigated, indicate that sites were dominated by Madoqua
(Dik-dik), lagomorpha, large to medium-sized bovids, giraffes,
carnivores, primates and elephants. The diversity of animals
represented in the footprint tuffs suggests that the landscape
of Pliocene Laetoli consisted of a variety of microhabitats.
Therefore, our results are consistent with inferences that Laetoli
was a mosaic landscape during the Pliocene (Musiba and
Magori, 2006; Harrison, 2005; Andrew, 1989).
This study demonstrates the potential of ichnofossils as eco-
logical indicators of time-averaged depositional environments
in paleontological sites. However, our results are limited by
our small sample size. Because of budget and time limitations,
we studied only six out of the 18 localities that contain animal
trackways reported by Mary Leakey and her co-workers in 1987.
Furthermore, the inherent assumption that Laetoli animals 3.8
million years ago may have preferred similar habitats to their
extant relatives may be open to question and further testing.
While we are aware of these limitations, the data provided
can enhance future studies of Laetoli’s paleoenvironments.
Future research is planned that will include analysis of bovid
limb elements and a more detailed analysis of the animal
tracks. The importance of Laetoli to our understanding of
human evolution compels consideration of the ichnofossil
record in reconstructions of its paleoenvironments. Also, the
ecological significance of the animal trackways warrants a major
conservation effort to preserve them.
We would like to thank our academic institutions for
facilitating the establishment of the field school and the semester
abroad program at Laetoli. We are particularly indebted to
our students who shared their field school experiences with
us, the Associated Colleges of the Midwest (ACM) for the
establishment of the Semester Abroad Program in Tanzania,
Dr. E.B. Chausi and the entire personnel at Ngorongoro
Conservation Area Authority (NCAA) for their understanding
of the scientific needs for such programs in the NCAA, and for
their invaluable time and contribution that shaped this study.
We also thank our Tanzanian colleagues, Ferdinand
Mizambwa, Felix Ndunguru, Godfrey Ole Moita, Said Killindo
and O.S. Kileo for comments, their help in field logistics and
field assistance during our summer field work at Laetoli. We are
very grateful to the Tanzanian Ministry of Tourism and Natural
resources and the Antiquities Department. We are also very
indebted to Mr. Donatius Kamamba for issuing the Antiquities
excavation license and to the doctor in-charge at Endulen
Hospital with his entire staff for the health related support to
our students and field staff.
We extend our thanks to the Lutheran Bishop of Dodoma,
Rev. Dr. Peter L. Mwamasika and Dr. Abel Nkini for their en-
couragement and transport-related logistic supports in Tanzania.
This project was funded by the Werner Gren Foundation (Gr.
5980), the Connecticut State University System, the ACM, the
National Geographic Society and a generous grant from Dr.
Melissa K. Stoller to Dr. Charles Musiba. We thank the editors
Downloaded by [Auraria Library] at 15:27 10 August 2014
of this volume and the two anonymous reviewers for providing
insightful comments from which the quality of this manuscript
benefited tremendously.
Agnew, N. and Demas, M. 1998. Preserving the Laetoli footprints. Scientific
American, 279(3):44–55.
Agnew, N. and Demas, M. 2000. Laetoli project: Report on the 1996–1997
field seasons and the Olduvai Museum exhibition. The Getty Conservation
Andrew, P. J. 1989. Palaeoecology of Laetoli. Journal of Human Evolution,
Bonnefille, R. and Riollet, G. 1987. Palynological spectra from the Upper Laetoli
beds. In Leakey, M. D. and Harris, J. M. (eds.), Laetoli: A Pliocene site in
northern Tanzania. Clarendon Press, Oxford: p. 52–61.
Butzer, K. W. 1974. Paleoecology of South African australopithecines: Taung
revisited. Current Anthropology, 15:376–382.
Coffing, E., Feibel, C., Leakey, M., and Walker, A. 1994. Four-million-year-old
hominids from east Lake Turkana, Kenya. American Journal of Physical
Anthropology, 93(1):55–65.
Coppens, Y. F., Howell, C., Isaac, G. L. I., and Leakey, R. E. F. 1976. Earliest
man and environments in the Lake Rudolf basin: Introduction. In Coppens,
Y. F., Howells, C., Isaac, G. L.I., and Leakey, R. E. F. (eds.), Earliest man
and environments in the Lake Rudolf basin. University of Chicago Press,
Chicago: 615p.
Day, M. H. and Magori, C. C. 1980. A new fossil hominid skull (L.H. 18) from
the Ngaloba beds, Laetoli, northern Tanzania. Nature, 284:55–56.
Day, M. H. and Wickens, E. H. 1980. Laetoli Pliocene hominid footprints and
bipedalism. Nature, 286:385–387.
Demas, M., Agnew, N., Waane, S., Podany, J., Bass, A., and Kamamba,
D. 1996. Preservation of the Laetoli hominid trackway in Tanzania.
In Roy, A. and Smith, P. (eds.), Preprints of the contributions to the
Copenhagen Congress, August 26–30, Archaeological Conservation and
its Consequences. International Institute for Conservation of Historic and
Artistic Works, London: p. 38–42.
Denys, C. 1987. Rodentia and lagomorpha. In Leakey,M.D.andHarris,J.
M. (eds.), Laetoli: A Pliocene site in northern Tanzania. Clarendon Press,
Oxford: p. 118–170.
Drake, R. and Curtis, G. H. 1987. Geochronology of the Laetoli fossil localities.
In Leakey,M.D.andHarris,J.M.(eds.),Laetoli: A Pliocene site in northern
Tanzania. Clarendon Press, Oxford: p. 48–52.
Feibel, C. S., Brown, F. H., and McDougall, I. 1989. Stratigraphic context
of the fossil hominids from the Omo group deposits: Northern Turkana
basin, Kenya and Ethiopia. American Journal of Physical Anthropology,
Feibel, C. S., Agnew, N., Latimer, B., Demas, M., Marshall, F., Waane, S. A. C.,
and Schmid, P. 1995. The Laetoli hominid prints—a preliminary report
on the conservation and scientific restudy. Evolutionary Anthropology,
Gentry, A. W. 1981. Pliocene and Pleistocene bovidae in Africa. In Beden,
M., Eisennmann, V., Geraads, D., and Gu
erin, C. (eds.), L’environnement
des hominid
es au Plio-Pl
ene. Fondation Singer Polignac, Paris: p.
Gentry, A. W. 1987. Pliocene bovidae from Laetoli. In Leakey, M. D. and Harris,
J. M. (eds.), Laetoli, A Pliocene site in northern Tanzania. Clarendon Press,
Oxford: p. 378–408.
Harris, J. M. 1985. Age and paleoecology of the Upper Laetolil beds, Laetoli,
Tanzania. In Ancestors: The hard evidence. Alan Riss, Inc., New York, p.
Harrison, T. 2005. Fossil bird eggs from the Pliocene of Laetoli, Tanzania:
Their taxonomic and paleoecological relationships. Journal of African Earth
Sciences, 41:289–302.
Hay, R. L. 1978. Melilitite-Carbonatite tuffs in the Laetoli Beds of Tanzania.
Contributions to Mineralogy and Petrology, 67:357–367.
Hay, R. L. 1980. Paleoenvironment of the Laetoli beds, northern Tanzania. In
Rapp, G. and Vondra, C. F. (eds.), Hominid sites: Their geologic settings.
AAAS Selected Symposium 63. Westview Press Inc., Boulder: p. 7–24.
Hay, R. L. and Leakey, M. D. 1982. The fossil footprints of Laetoli. Scientific
American, 246:50–57.
Hay, R. L. 1987. Geology of the Laetoli area. In Leakey,M.D.andHarris,J.
M. (eds.), Laetoli: A Pliocene site in northern Tanzania. Clarendon Press,
Oxford: p. 23–47.
Johanson, D. and Edey, M. A. 1981. Lucy, the beginnings of humankind. Simon
and Schuster, New York, 409p.
Kent, P. E. 1941. The recent history and Pleistocene deposits of the plateau north
Lake Eyasi. Tanganyika. Geological Magazine, 78:173–184.
Kingdon, J. 1982. East African mammals, Vol. III. University of Chicago Press,
Chicago, IL: 358p.
Kingdon, J. 1997. The kingdom field guide to African mammals. Academic
Press, London: 450 pp.
Kyauka, P. S. 1994. Developmental patterns of the earliest hominids: Amorpho-
logical perspective. In Corruccini, R. S. and Ciochon, R. L. (eds.), Integrative
paths to the past: Paleoanthropological advances in honor of F. Clark Howell.
Prentice Hall, Englewood Cliffs, NJ: p. 229–250.
Leakey, M. D. 1981. Tracks and tools. Philosophical Transactions Royal Society
London, B 292:95–102.
Leakey, M. D. 1987. The hominid footprints: Introduction. In Leakey,M.D.and
Harris, J. M. (eds.), Laetoli: A Pliocene site in northern Tanzania. Clarendon
Press, Oxford: p. 490–496.
Leakey, M. D., Hay, R. L., Curtis, G. H., Drake, R. E., Jackes, M. K., and White,
T. D. 1976. Fossil hominids from the Laetolil beds. Nature, 262:460–466.
Leakey, M. D. and Hay, R. L. 1979. Pliocene footprints in the Laetolil beds at
Laetoli, northern Tanzania. Nature, 278:317–323.
Leakey, M. D., Beden, M., Gu
erin, C., Renders, E. M., and Sondaar, P. 1987.
Animal prints and trails. In Leakey,M.D.andHarris,J.M.(eds.),Laetoli:
A Pliocene site in northern Tanzania, Clarendon Press, Oxford: p. 451–489.
Manega, P. C. 1993. Geochronology, geochemistry and isotopic study of the
Plio-Pleistocene hominid sites and the Ngorongoro volcanic highland in
northern Tanzania. Ph.D. thesis, University of Colorado, Boulder, Colorado.
Musiba, C. M. 1999. Laetoli Pliocene Paleoecology: A Reanalysis via
morphological and behavioral approaches. Ph.D. thesis, University of
Musiba, C. M. and Magori, C. C. 2006. Functional morphology, taphonomy
and paleoecological context of the Upper Laetolil beds (localities 8 & 9),
Laetoli in northern Tanzania. African Genesis: A Symposium on Hominid
Evolution in South Africa, January 8–14, University of the Witwatersrand
Medical School, Johannesburg, South Africa.
Ndessokia, P. N. S. 1990. The mammalian fauna and archaeology of the
Ndolanya and Olpiro beds, Laetoli, Tanzania. Ph.D. thesis, University of
California, Berkeley.
Stuart, C. and Stuart, T. 1994. A field guide to the tracks and signs of southern
and East African wildlife. Struik Publishers, Cape Town: 310 pp.
Tuttle, R. H. 1987. Kinesiological inferences and evolutionary implications from
Laetoli bipedal trails G-1, G-2/3, and A. In Leakey,M.D.andHarris,J.
M. (eds.), Laetoli: A Pliocene site in northern Tanzania. Clarendon Press,
Oxford: p. 503–523.
Tuttle, R. H. 1992. Footprints and gaits of bipedal apes, bears, and barefoot
people: Perspective on Pliocene tracks. In Matano, S., Tuttle, R. H., Ishida,
H., and Goodman, M. (eds.), Topics in primatology, vol. 3. University of
Tokyo Press, Tokyo: p. 221–242.
Verdcourt, B. 1987. Mollusca from the Laetolil and Upper Ndolanya beds. In
Leakey,M.D.andHarris,J.M.(eds.),Laetoli: A Pliocene site in northern
Tanzania. Clarendon Press, Oxford: p. 438–450.
Vrba, E. S. 1975. Some evidence of chronology and paleoecology of
Sterkfontein, Swartkrans and Kromdraai from the fossil Bovidae. Nature,
Downloaded by [Auraria Library] at 15:27 10 August 2014
Vrba, E. S. 1985a. Environment and evolution: Alternative causes of the
temporal distribution of evolutionary events. South African Journal of
Science, 81:229–236.
Vrba, E. S. 1985b. African bovidae: Evolutionary events since
the Miocene. South African Journal of Science, 81:263–
Vrba, E. S. 1995. The fossil record of African antelopes (Mammalia, Bovidae)
in relation to human evolution and paleoclimates. In Vrba, E. S., Denton, G.
H., Partridge, T. C., and Burckle, L. H. (eds.), Paleoclimate and evolution,
with emphasis on human origins. Yale University Press, New Haven, CT: p.
Vrba, E. S. 2000. Major features of Neogene mammalian evolution in Africa.
In Partridge, T. C. and Maud, R. R. (eds.), The cenozoic of southern Africa.
Oxford University Press, Oxford: p. 277–304.
White, T. D. 1977. New fossil hominids from Laetoli, Tanzania. American
Journal of Physical Anthropology, 46:197–230.
White, T. D. 1980. Additional fossil hominids from Laetoli, Tanzania. American
Journal of Physical Anthropology, 53:487–504.
WoldeGabriel, G., White, T. D., Suwa, G. Renne, P., de Heinzelin, J., Hart,
W. K., and Helken, G. 1994. Ecological and temporal placement of early
Pliocene T. Nature, 371:330–333.
Wood, B. and Collard, M. 1999. The human genus. Science, 284:65–71.
Downloaded by [Auraria Library] at 15:27 10 August 2014
... Полевые исследования, проведенные в последние годы (Musiba et al., 2008(Musiba et al., , 2020Getti…, 2001Getti…, , 2011Zaitsev et al., 2019Zaitsev et al., , 2020, показывают, что происходит постепенное разрушение туфов Лаетоли, а также разнообразных отпечатков в них вследствие физического и химического выветривания и необходимы меры для разработки технологии по консервации и сохранению отпечатков Australopithecus afarensis. В данной статье мы рассматриванием возможные физико-химические условия, при которых происходит разрушение первичных минералов туфов, а также образование вторичных минералов в этих ассоциациях. ...
... Наиболее изученным является маркирующий туф № 7 (Hay, 1987;Barker and Milliken, 2008;McHenry, 2011;Zaitsev et al., 2011Zaitsev et al., , 2015Zaitsev et al., , 2019Zaitsev et al., , 2020. Это обусловлено тем, что именно в нем были обнаружены многочисленные отпечатки следов древних животных, а также отпечатки ступней Australopithecus afarensis (Leakey and Hay, 1979;Musiba et al., 2008;Zaitsev et al., 2020). В разрезе туфа выделяются четыре слоя общей мощностью до 50 см, которые различаются по цвету, текстуре, структуре и соотношению первичных и вторичных минералов -авгит-биотитовый, слоистый, белый и серый туфы (рис. ...
... Физическое и химическое изменение туфов Лаетоли происходило как в момент формирования пород, так и в настоящее время. На это указывают результаты исследований, выполненных Ч. Мусибой с соавторами (Musiba et al., 2008), специалистами института Гетти (Getty…, 2001(Getty…, , 2011, и наши данные, полученные в 2016 г. (Musiba et al., 2020). Сравнение фотографий отпечатков, сделанных в различные годы, показывает существенные изменения в морфологии отпечатков; мы полагаем, что, с одной стороны, происходит физическое разрушение кальцита вследствие высокого статического давления со стороны консервационного кургана (см. ...
The Laetoli area in northern Tanzania is an important palaeo-anthropological site, where the oldest footprints of Australopithecus afarensis reside. Aeolian tuffs are the major rock type at Laetoli and they are divided into Lower and Upper Laetolil Beds that were deposited at an interval of 4.36 and 3.63 million years. The Upper Laetolil Beds contain eight layers of air-fall tuffs known as marker tuffs. The Australopithecus afarensis footprints are observed on the surface of the white tuff, which is a part of the Upper Laetolil marker tuff 7, also known as the “Footprint Tuff.” The interpolated age of the marker tuff 7 is 3.66 million years. Two mineral assemblages are distinguished in the Upper Laetolil marker tuffs. The first assemblage consists of primary tuff minerals and includes clinopyroxene (diopside, augite, aegirine-augite), nepheline, melilite (åkermanite and alumoåkermanite), garnet (andradite and schorlomite), magnetite, and others. The second mineral assemblage consists of secondary minerals, montmorillonite, calcite, and phillipsite. They were formed during replacement of the primary minerals, volcanic glass, and ash cementation. Thermodynamic calculations show that the major primary tuff minerals (melilite and nepheline) are stable at variable sodium activity and pH values. Replacement of melilite and nepheline by montmorillonite is caused by a decrease of sodium activity in slightly alkaline, neutral and acidic conditions (рН < 10). Montmorillonite is not present in the altered nephelinitic tuff of the Sadiman volcano (which is considered as a source of the Laetolil Beds) where kaolinite is the major secondary mineral. This is explained by the difference in H2O fugacity with higher lgfH2O values in Sadiman and lower values in Laetoli. Relationships between primary and secondary tuffs minerals on the lgaHCO3 vs pH plot suggest mineral transformation within the Laetolil Beds in slightly acid and neutral con- ditions (рН = 5–7) compared with more alkaline conditions at Sadiman (рН > 10).
... There are lots of examples where a trackmaker is inferred without such description (e.g. Musiba et al. 2008). In describing and/or measuring a track of whatever origin, we have three main independent properties to consider. ...
... Mammal, bird, and insect prints and trails were identified by Mary Leakey and collaborators in 18 sites (labelled from A to R) out of 33 total palaeontological 134 M. Cherin et al. localities in the Laetoli area (Leakey 1987;Harrison and Kweka 2011;Musiba et al. 2008). The so-called Footprint Tuff, which corresponds to the lower part of Tuff 7 in the Upper Laetolil Beds' stratigraphic sequence, hosts at least ten sublevels in which footprints are found (Hay 1987). ...
... The site consists of 18 large ichnological surfaces Leakey and Harris 1987). Thousands of fossilized animal tracks have been found there over the years (Leakey and Harris 1987;Musiba et al. 2008). In the late 1970s, a trail of hominin footprints was found at Laetoli's Site G. ...
... Unfortunately, many important ichnological sites are in countries that lack the necessary skills and resources to implement conservation projects or to build adequate protective or museum structures. Indeed, the scientific community has repeatedly sounded the alarm about the critical state in which the Laetoli footprints have been allowed to fall (Dalton 2008;Musiba et al. 2008). ...
Full-text available
We currently have little archaeological and historical data about mining areas of North Africa that were exploited in pre-Roman times, and even the rare ancient proofs can be mostly dated to Roman period. However, it was the searching for new metals that brought Phoenician to the western expansion: North Africa was one of the richest Mediterranean areas and we believe that such a small amount of information about this region has to be connected to the lack of dedicated studies. Every time that the scientific interest focuses on this important part of the economic and social life in the ancient world (as it was already made in Spain and Sardinia), new data emerges and offers a bigger historical background. Current researches of ‘Researches about Phoenicians in North Africa: archaeology, numismatic and economic history’ aims, first of all, to define historical and technological background of all the metal production cycle, the exploitation, the manipulation techniques and the resources management of mining areas in North Africa, Morocco and Algeria in particular
... While the sites described above offer fascinating insights into the environments and animal communities directly associated with fossil hominins, the potential for trace fossils to inform paleoenvironmental reconstructions remains underexplored. In addition to the studies reviewed here, Musiba and colleagues 91 ...
Hominin footprints have not traditionally played prominent roles in paleoanthropological studies, aside from the famous 3.66 Ma footprints discovered at Laetoli, Tanzania in the late 1970s. This contrasts with the importance of trace fossils (ichnology) in the broader field of paleontology. Lack of attention to hominin footprints can probably be explained by perceptions that these are exceptionally rare and "curiosities" rather than sources of data that yield insights on par with skeletal fossils or artifacts. In recent years, however, discoveries of hominin footprints have surged in frequency, shining important new light on anatomy, locomotion, behaviors, and environments from a wide variety of times and places. Here, we discuss why these data are often overlooked and consider whether they are as "rare" as previously assumed. We review new ways footprint data are being used to address questions about hominin paleobiology, and we outline key opportunities for future research in hominin ichnology.
... The only other southern African record that we are aware of is from Namibia, where Bennett et al. (2010) describe Holocene elephant tracks in the Kuiseb Delta. Elsewhere in Africa, Pliocene elephant tracks have been noted in Laetoli, Tanzania (Musiba et al., 2008), and Pleistocene elephant tracks were found in Ileret, Kenya (Roach et al., 2016). ...
Full-text available
Aeolianites and cemented foreshore deposits on South Africa's Cape south coast have the capacity to record and preserve events that transpired on them when they were composed of unconsolidated sand. Thirty-five Pleistocene elephant tracksites have been identified along this coastline. This abundance of sites along what was the margin of the vast Palaeo-Agulhas Plain allows for an appreciation of the forms that elephant tracks and traces can take in the context of the global proboscidean track record. They point to a significant regional elephant presence from Marine Isotope Stage (MIS) 11 (~400 ka) through MIS 5 (~130–80 ka) to MIS 3 (~35 ka) and also indicate repeated use of certain dune areas. They buttress Holocene and historical evidence that elephants made use of open areas in the region, and that the remaining “Knysna elephants” retreated into dense afrotemperate forest for protection in recent centuries. Analogies can be drawn between Pleistocene elephant tracks and Mesozoic dinosaur tracks, and some of the Cape south coast elephant tracks are among the largest Cenozoic (and hence, Quaternary) tracks ever to be described. A newly identified tracksite in this area may provide the first reported evidence of elephant trunk-drag impressions.
... Mammal, bird, and insect prints and trails were identified by Mary Leakey and collaborators in 18 sites (labelled from A to R) out of 33 total palaeontological localities in the Laetoli area (Leakey 1987;Harrison and Kweka 2011;Musiba et al. 2008). The so-called Footprint Tuff, which corresponds to the lower part of Tuff 7 in the Upper Laetolil Beds' stratigraphic sequence, hosts at least ten sublevels in which footprints are found (Hay 1987). ...
Full-text available
The Romanian karst hosts numerous caves and shelters that over time provided remarkable archaeological and anthropological vestiges. Altogether they show that humans must have entered caves in Romania at least as early as 170,000 years ago. However, ancient human footprints are very rare in the fossil record of East-Central Europe, with only two known locations in the Apuseni Mountains of western Romania. Vârtop Cave site originally preserved three fossil footprints made about 67,800 years ago by a Homo neanderthalensis , whereas Ciur Izbuc Cave was probably home of early H. sapiens that left almost 400 footprints (interspersed with spoors of cave bears), which were indirectly dated to be younger than ~36,500 years.
... There are lots of examples where a trackmaker is inferred without such description (e.g. Musiba et al. 2008). In describing and/or measuring a track of whatever origin, we have three main independent properties to consider. ...
Full-text available
Animal footprints are preserved in the archaeological record with greater frequency than perhaps previously assumed. This assertion is supported by a rapid increase in the number of discoveries in recent years. The analysis of such trace fossils is now being undertaken with an increasing sophistication, and a methodological revolution is afoot linked to the routine deployment of 3D digital capture. Much of this development has in recent years been driven by palaeontologists, yet archaeologists are just as likely to encounter footprints in excavations. It is therefore timely to review some of the key methodological developments and to focus attention on the inferences that can and, crucially, cannot be justifiably made from fossil footprints with specific reference to human tracks.
... localities in the Laetoli area (Leakey 1987;Harrison and Kweka 2011;Musiba et al. 2008). The so-called Footprint Tuff, which corresponds to the lower part of Tuff 7 in the Upper Laetolil Beds' stratigraphic sequence, hosts at least ten sublevels in which footprints are found (Hay 1987). ...
Full-text available
Fossil footprints are very useful palaeontological tools. Their features can help to identify their makers and also to infer biological as well as behavioural information. Nearly all the hominin tracks discovered so far are attributed to species of the genus Homo. The only exception is represented by the trackways found in the late 1970s at Laetoli, which are thought to have been made by three Australopithecus afarensis individuals about 3.66 million years ago. We have unearthed and described the footprints of two more individuals at Laetoli, who were moving on the same surface, in the same direction, and probably in the same timespan as the three found in the 1970s, apparently all belonging to a single herd of bipedal hominins walking from south to north. The estimated stature of one of the new individuals (about 1.65 m) exceeds those previously published for Au. afarensis. This evidence supports the existence of marked morphological variation within the species. Considering the bipedal footprints found at Laetoli as a whole, we can hypothesize that the tallest individual may have been the dominant male, the others smaller females and juveniles. Thus, considerable differences may have existed between sexes in these human ancestors, similar to modern gorillas.
Humans evolved in the dynamic landscapes of Africa under conditions of pronounced climatic, geological and environmental change during the past 7 million years. This book brings together detailed records of the paleontological and archaeological sites in Africa that provide the basic evidence for understanding the environments in which we evolved. Chapters cover specific sites, with comprehensive accounts of their geology, paleontology, paleobotany, and their ecological significance for our evolution. Other chapters provide important regional syntheses of past ecological conditions. This book is unique in merging a broad geographic scope (all of Africa) and deep time framework (the past 7 million years) in discussing the geological context and paleontological records of our evolution and that of organisms that evolved alongside our ancestors. It will offer important insights to anyone interested in human evolution, including researchers and graduate students in paleontology, archaeology, anthropology and geology.
Full-text available
Bipedal trackways discovered in 1978 at Laetoli site G, Tanzania and dated to 3.66 million years ago are widely accepted as the oldest unequivocal evidence of obligate bipedalism in the human lineage 1–3 . Another trackway discovered two years earlier at nearby site A was partially excavated and attributed to a hominin, but curious affinities with bears (ursids) marginalized its importance to the paleoanthropological community, and the location of these footprints fell into obscurity 3–5 . In 2019, we located, excavated and cleaned the site A trackway, producing a digital archive using 3D photogrammetry and laser scanning. Here we compare the footprints at this site with those of American black bears, chimpanzees and humans, and we show that they resemble those of hominins more than ursids. In fact, the narrow step width corroborates the original interpretation of a small, cross-stepping bipedal hominin. However, the inferred foot proportions, gait parameters and 3D morphologies of footprints at site A are readily distinguished from those at site G, indicating that a minimum of two hominin taxa with different feet and gaits coexisted at Laetoli.
Reviews previous notions of what might cause speciation and extinction, and hypotheses that have predicted temporal patterns of evolutionary events. Discusses a new postulate relating to the 'turnover-pulse' (a concentration of turnover events against the time scale), arguing that global climatic forcing of speciation rhythms has been of major importance to the diversification of mammals on the huge continental landmass of Africa in the Neogene. The turnover-phase hypothesis uses concepts of evolutionary conservation, species' habitat-specificity and vicariance. -P.J.Jarvis
Bovid (essentially antelope) biostratigraphy and phylogeny indicate a widespread environmental change in Africa near 2.5 Myr. Morphological and ecological analyses of the taxa lead to the inference that across sub-Saharan Africa a spreading of open grassland, probably caused by global reduction in temperature and associated changes in rainfall, occurred at around this time. -from Author
The Laetoli hominid footprints in northwestern Tanzania are of major importance in understanding the evolution of humankind. The tracks are 3.6 million years old and pre-date the development of tool-making by one million years, providing clear evidence of the precedence of bipedalism over evolution of the brain. The site was excavated in the late 1970s by Mary Leakey, documented and reburied. In subsequent years, re-vegetation occurred and damage from root growth resulted. Of particular interest in relation to reburial as a preservation strategy is the fact that serious damage occurred to a unique scientific and cultural site because subsequent monitoring and maintenance of the reburial were not carried out. This paper presents an overview of the strategy developed between 1992 and 1994 for the preservation of the trackway, which takes into consideration the many and complex requirements of preserving outdoor sites, and describes the re-excavation, conservation, documentation and reburial of the southern 10m of the trackway completed in the 1995 field season.
Animal tracks and hominid footprints have been discovered in the Laetolil Beds of Tanzania. The volcanic-ash deposits were formed between 3.5 and 3.8 million years ago by the eruption of the volcano sadiman. The Laetolil Beds are north of Lake Eyasi and southeast of Lake Victoria. Early hominid remains have been found near the area. The footprint tuff lies near the top of the fossil-bearing strata. Thousands of animal tracks have been discovered in the ash deposits. The savanna supported an abundant and diverse animal population. The ash buried animal bones, teeth, eggs and dung. Fossilized twigs, shrubs, thorns and leaves provide information on the plants of the savanna. The hominid tracks are proof that they walked fully upright. No stone tools have been found in the Laetolil Beds.-F.McElhoe Jr
The environmental context and stratigraphic position of the first australopithecine discovery, at Taung, South Africa, need to be reconsidered, since the Taung juvenile, as Australopithecus africanus, is the type specimen of the gracile lineage of Plio-Pleistocene hominids. Recent study of the site and its regional setting shows that two generalizations widely held today are either incorrect or dubious. In particular, the Taung hominid is found in deposits that clearly indicate a subhumid or humid environment and not a semidesert, thus questioning the original basis for hypotheses concerning an ecological differentiation of gracile and robust australopithecines. Furthermore, the Taung specimen relates to a younger geomorphologic cycle than postulated by F. E. Peabody, and this cycle appears to immediately predate the Middle Pleistocene "Younger Gravels" of the Vaal River. This and other, faunal criteria suggest that Taung is contemporary with or even younger than Swartkrans and Kromdraai rather than broad...