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Journal of Human Evolution 59 (2010) 123e132

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Journal of Human Evolution


journal homepage: www.elsevier.com/locate/jhevol

New evidence for a 67,000-year-old human presence at Callao Cave,


Luzon, Philippines
Armand Salvador Mijares a, *, Florent Dtroit b, Philip Piper a, Rainer Grn c, Peter Bellwood d,
Maxime Aubert c, Guillaume Champion b, Nida Cuevas e, Alexandra De Leon e, Eusebio Dizon e
a

Archaeological Studies Program, Palma Hall, University of the Philippines, Diliman, Quezon City 1101, Philippines
UMR 7194, CNRS, Dpartement de Prhistoire du Musum national dhistoire naturelle, 57, rue Cuvier, 75005 Paris, France
Research School of Earth Sciences, Bldg 61 Mills Road The Australian National University, Canberra ACT 0200, Australia
d
School of Archaeology and Anthropology, AD Hope Building, The Australian National University, Canberra ACT 0200, Australia
e
Archaeology Division, National Museum of the Philippines, P Burgos Ave., Manila, Philippines
b
c

a r t i c l e i n f o

a b s t r a c t

Article history:
Received 24 August 2009
Accepted 22 April 2010

Documentation of early human migrations through Island Southeast Asia and Wallacea en route to
Australia has always been problematic due to a lack of well-dated human skeletal remains. The best
known modern humans are from Niah Cave in Borneo (40e42 ka), and from Tabon Cave on the island of
Palawan, southwest Philippines (47  11 ka). The discovery of Homo oresiensis on the island of Flores in
eastern Indonesia has also highlighted the possibilities of identifying new hominin species on islands in
the region. Here, we report the discovery of a human third metatarsal from Callao Cave in northern
Luzon. Direct dating of the specimen using U-series ablation has provided a minimum age estimate of
66.7  1 ka, making it the oldest known human fossil in the Philippines. Its morphological features, as
well as size and shape characteristics, indicate that the Callao metatarsal denitely belongs to the genus
Homo. Morphometric analysis of the Callao metatarsal indicates that it has a gracile structure, close to
that observed in other small-bodied Homo sapiens. Interestingly, the Callao metatarsal also falls within
the morphological and size ranges of Homo habilis and H. oresiensis. Identifying whether the metatarsal
represents the earliest record of H. sapiens so far recorded anywhere east of Wallaces Line requires
further archaeological research, but its presence on the isolated island of Luzon over 65,000 years ago
further demonstrates the abilities of humans to make open ocean crossings in the Late Pleistocene.
2010 Elsevier Ltd. All rights reserved.

Keywords:
Cave faunas
Hominin dispersal
Southeast Asia
U-series dating

Introduction
Hominin movement into Island Southeast Asia has always been
problematic due to the lack of well-dated human remains. The
humid tropical environment of Island Southeast Asia contributes to
the problems of bone preservation. Early modern human remains
have, however, been recovered in Niah Cave in Sarawak, Malaysian
Borneo (Harrisson, 1975; Barker et al., 2002), dating to 42 ka
(Barker et al., 2007), and from Tabon Cave in Palawan (Fox, 1970;
Dizon et al., 2002), dating to 47  10/11 ka (Dtroit et al., 2004).
Borneo is located on the Sunda shelf and was possibly joined by dry
land to Sumatra, Java and Peninsular Malaysia during periods of
lowered sea level in the Pleistocene. The island of Palawan may
have been intermittently attached to northeastern Borneo when

* Corresponding author.
E-mail address: armand.mijares@up.edu.ph (A.S. Mijares).
0047-2484/$ e see front matter 2010 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jhevol.2010.04.008

sea levels reached their minima during the most extreme climatic
phases. Thus, migrating human populations could have reached
both islands without necessarily requiring a sea crossing. To reach
the rest of the Philippine archipelago and other islands in the
Wallacean group (e.g., Sulawesi, Flores, Timor) that were never
attached to either mainland Asia or Australasia (Sahul), open sea
crossings were required. The Lake Mungo remains from Australia
dating to 40  2 ka (Bowler et al., 2003) are evidence that modern
humans were capable of making very early sea crossings. Homo
oresiensis, discovered on the islands of Flores, Indonesia, is another
hominin that managed to cross the Wallace line. While its remains
are only dated to 18e38 ka (Morwood et al., 2004), Flores also has
stone artifact assemblages suggesting that a hominin of unknown
afnity reached the island more than 800 ka years ago (Brumm
et al., 2006). Our recent excavations (2007) in Callao Cave (Fig. 1)
have produced what is probably one of the earliest hominin fossils
east of Wallaces Line, from the island of Luzon, northern
Philippines.

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A.S. Mijares et al. / Journal of Human Evolution 59 (2010) 123e132

Figure 1. The Location of Callao Cave and the 2003 excavation.

Materials and methods


The excavation of Callao Cave
In 2003, archaeological excavations under the direction of
A. Mijares at Callao Cave in Peablanca, Cagayan Valley, northern
Luzon (Fig. 1), were designed to investigate the regional transition
from Paleolithic hunting and gathering to the introduction of
farming during the MideLate Holocene. At the commencement,
there was no intention of exploring the earlier history of the cave,
but the excavations exposed an Upper Pleistocene layer 130 cm
below present ground level that contained chert ake tools, burnt
animal bones and a hearth feature. A radiocarbon determination of
25 968  374 (uncalibrated) BP (Mijares, 2005, 2007) made this the
oldest known human occupation in the main Philippine archipelago (excluding Palawan) and demonstrated that there was
Paleolithic occupation in the cave entrance. To investigate this
further, a joint partnership between the University of the
Philippines Archaeological Studies Program, the National Museum
of the Philippines, and the Australian National University was
established to continue excavation in Callao Cave in 2007.
The 2007 excavations started at 130 cm below surface, the level
at which the 2003 excavation had ended (Fig. 2). Procedures
followed the 2003 system of excavating in 5 cm spits and recording
each archaeological nd three dimensionally by spit and stratigraphic layer. The excavation was reduced in size from the 2003
excavations, and only Square 1 in the north (2  2 m) and the
northern portion of Square 2 (2  1 m) were excavated below
130 cm. Since water for wet sieving was not available in the cave,
most sediment was dry sieved through a 4 mm mesh. In addition,
samples were also collected for otation.
At a depth of 160 cm below the surface (BS), the excavation area
was further reduced and only the southern end of Square 1 and the
northern end of Square 2 were excavated. Between 160 cm and
250 cm, BS archaeological remains were scarce, with just a ake
tool, chert core and deer bones recovered from Layer 11, and several
poorly preserved deer bones, including a scapula, the proximal end
of a humerus and several fragments of antler identied in Layer 12.
There was no evidence of human occupation similar to that
observed in the Upper Pleistocene Layer 8.

In Layer 13 (255e265 cm BS), the bones and teeth of deer


became more abundant, and in Layer 14 (270e295 cm BS), the team
encountered cemented sediment (carbonized breccia) containing
a relatively dense concentration of animal bone. In this layer, at
a depth of 275 cm below the cave surface, we found the human
third metatarsal (MT3) bone, which is the subject of this paper.
Beyond 300 cm depth, the excavation unit became too conned
between the cave wall and a large stalagmite for excavation to
continue.

Figure 2. Stratigraphic prole of Square 2, Callao Cave. Layer 14 is the breccia that
contained the human metatarsal (Callao MT3).

A.S. Mijares et al. / Journal of Human Evolution 59 (2010) 123e132

Dating
Three osseous samples from the breccia were selected for
dating. Two cervid teeth (Callao 1 and Callao 2) recovered from 275
to 295 cm below present ground surface were dated by Electron
Spin Resonance (ESR) and Uranium Series (U-series) analysis. Laser
ablation U and U-series analysis was carried out at by R. Grn at the
Australian National University (see Eggins et al., 2003, 2005 for the
experimental set-up and Grn et al., 2005, 2006, 2008 for previous
applications). The ESR dating of the teeth followed procedures
routinely applied in the ANU ESR dating laboratory (Grn et al.,
2001). For ESR dose analysis, the enamel was powdered and
aliquots were successively irradiated in ten steps to 372 Gy.
Radiation doses were monitored with alanine dosimeters and
evaluated against a calibrated dosimeter set provided by A. Wieser,
Messtechnik, Mnchen. Dose values were determined by partial
spectrum tting (Grn, 2002). The external dose rate was determined from a single representative sample from the breccia. The
U-concentration in the dentine of the two cervid teeth had only
small variations along the laser track, 71.6  8.2 ppm U for Callao 1
and 100  5 ppm U for Callao 2, as usually observed in dentine.
In order to obtain an age estimate for the hominin metatarsal,
the Callao MT3 was also directly analyzed. A diamond wire saw was
used to cut the bone 2 mm away from the break described below.
Four laser ablation scans were then recorded on the cross-section
(Fig. 3).
Preservation, conservation and cleaning of the Callao MT3
The Callao specimen with National Museum of the Philippines
accession number II-77-J3-7691 is a right third metatarsal (Fig. 4). It
is broken in two parts around the mid-shaft, but with the exception
of the distal head, which is broken, the bone is almost complete.
The missing head has not been identied among the other bones
recovered from the same layer. The overall state of preservation of
the bone is rather good, most being preserved under a thin
yellowish coat of calcium carbonate concretions. Several small
patches of hard and compact sediment are also attached to the
bone. Smaller scratches (which occurred presumably during excavation) are visible on the medial and lateral surfaces of the shaft
where the white cortex of the bone is exposed.
Both surfaces (proximal and distal) of the transverse break
around mid-shaft are covered by a very thin layer of carbonate
concretions, which indicates that the fracture is ancient. However,
it is neither oblique nor spiral, and has no helicoidal morphology.
No scars or peeling around the fracture are visible. This indicates
that breakage did not occur on fresh bone. This transverse break,

125

almost perpendicular to the long axis of the shaft, resulted from the
post depositional fracture of already dry bone (Lyman, 1994). The
two parts of the metatarsal join almost perfectly at the fracture, and
only the presence of a thin layer of concretion inhibits a perfect
retting.
Before cleaning the bone, a complete set of photographs, a 3D
surface scan and a mCT-scan were made for archival records and
future analysis. The object of cleaning the bone was to remove the
thickest patches of sediment in order to observe its shape and
morphological features. Whenever possible, the thin carbonated
coating was kept intact to avoid damaging the underlying surface,
which was cleaned using dental tools under a stereo-microscope.
Only puried water and a 50e50 mix of puried water and alcohol
were used to soften the compact sediment.
Except where noted, all descriptions and comparisons were
made after cleaning. Observations and measurements were made
directly on the original specimen and on the 3D surface scan, after
virtual retting of the proximal and distal portions of the bone at
the mid-shaft fracture.
Results
Faunal remains
From the 2007 eld season, the Callao Cave excavations
produced a late Pleistocene vertebrate assemblage of 807 fragments of bone. In Layer 11, just a few isolated and potentially
re-worked fragments of animal bone, including several skeletal
elements of deer, were recovered. There was no evidence of
anything that could be considered an in situ human-derived bone
accumulation and no evidence of butchery or lithics. Humans,
however, appear to have been present during this period of deposition, given the discovery of a terminal foot phalanx in Spit 36
(180 cm BS), with an interpolated date of 30 ka.
Bone concentrations increased below Spit 47 (Layer 12). Down
to Spit 53 (265 cm BS), all of the bone fragments were recovered
from clayey silt sediments and were very poorly preserved. Spit 54
contained numerous mid-dark brown skeletal elements, similar in
appearance to those recovered throughout the clayey silt deposits
above, as well as a few weathered and eroded fragments with
a creamy appearance, similar to those from the underlying breccia
(Layer 14). It is likely that, after the Layer 14 breccia had formed,
erosional processes caused some bones to be redistributed into the
accumulating clayey silt sediments above. Alternatively, it is
possible that the boundary between clayey silt and breccia has
migrated downwards as the breccia has decalcied, and that Spit 54
(270 cm BS) forms a transitional zone between breccia and clay.

Figure 3. Cross-section of the Callao metatarsal showing the scanned transects.

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A.S. Mijares et al. / Journal of Human Evolution 59 (2010) 123e132

Figure 4. The Callao right metatarsal III (II-77-J3-7691).

The breccia contains by far the greatest number of bone fragments (66%, or 533 fragments), indicating that bone survival was
good within the calcium carbonate precipitate. As a result, bones
from the breccia are in much better physical condition than the
pieces of bone recovered from the overlying clayey silts.
Surface modications consistent with subsurface weathering,
erosion and water-transport suggest that the bones were derived
from elsewhere in the cave system and were redeposited against
the wall, where they were incorporated into a carbonate-cemented
sandy silt loam breccia. The complete absence of any evidence
for bone-accumulating large carnivores and porcupines in the
Philippines (excluding Palawan), in addition to the presence of
three parallel cut-marks caused by a sharp implement on the
medial surface of a deer tibia shaft fragment (Fig. 5), indicates that
the bone assemblage was almost certainly derived from human
activity (Piper and Mijares, 2007). The lack of any lithics from the
1980 (Cuevas, 1980) and present excavations at the depth of the
breccias in Callao Cave could imply the use of organic tools. Alternatively, the differential transport of bones has caused spatial
separation of archaeological materials in the cave entrance.
Three large species of mammal were recorded in the Callao Cave
breccia: the native brown deer (Cervus mariannus), the Philippine
warty pig (Sus philippensis), and an extinct bovid evidenced by two
small tooth fragments (Piper and Mijares, 2007). Brown deer
constitute more than 90% of the identiable bone fragments. The
articular ends of long bones and metapodials are relatively
common in the assemblage. Loose maxillary and mandibular teeth
indicate that crania were originally present, and scapula, pelvic and
vertebral fragments indicate the deposition of axial elements as
well. This suggests that whole, or different skeletal elements of
various deer carcasses were possibly once brought into the cave on
occasion.

while Callao 2 did not yield a result (the closed system ESR age
estimate was younger than the U-series result). The discrepant
combined ESR/U-series results point to some reworking, as
conrmed by taphonomic examination of the bones from the
breccia.
It can be seen that the U-concentration varies greatly along the
proles (Fig. 6). We carried out U-series age estimates for all
concentration features to check whether delayed uptake or leaching has taken place, such as peaks and troughs (Fig. 6). We found
only a small number of results (seven of 69) where the 1  s
weighted means did not overlap. This is somewhat less than
expected from a normal statistical distribution and points to the
fact that our error bars are probably somewhat too large.
On the whole, there is no relationship between U-concentration
and age estimate (Fig. 7). This points to a rapid uptake and excludes
leaching along pores. Instead, as an indication for U-leaching, the
drops in U-concentration are probably caused by heterogeneities in
the bone structure, which had little inuence on the mode of
U-mobilization.
Single age estimates may be affected by a number of processes
rendering them inaccurate, and for this reason the U-series results
must be considered minimum estimates. This alone precludes any
claim for accuracy. In this regard, the interpretation of APNIAH1 in
Barker et al. (2007) has been used to cast doubt on the results of
U-Series dating at Niah. Barker et al. (2007, p. 253) write .a

Age determination
The cervid two teeth yielded U-series ages of 52  1.4 ka
(Callao 1) and 54.3  1.9 ka (Callao 2). Sub-sampling of small peaks
and troughs in the U-concentration had yielded U-series age estimates that were in the same statistical population as the mean
values. We could not identify any regions that may indicate delayed
U-uptake or leaching, compared with the average values. Thus, the
U-uptake must have taken place over a relatively short period of
time (Pike et al., 2002). However, as U-accumulation in bones may
be delayed after burial, any U-series age estimates have to be
considered minimum age estimates. ESR dating yielded a combined
ESR/U-series result (Grn et al., 1988) for Callao 1 of 66 11/9 ka,

Figure 5. Cut marks on the surface of a cervid bone.

A.S. Mijares et al. / Journal of Human Evolution 59 (2010) 123e132

127

Figure 6. Comparative diagrams of the four laser ablation scans on the Callao metatarsal.

uniform date prole [of APNIAH1] may also be reached after further
uptake or leaching of uranium, which would give an erroneous result,
so on its own the date should only be taken as provisional.. In the
context of MT3, we do not have to consider further U-uptake,
because this would only lower the calculated apparent U-series

estimates. The question is whether our results could seriously


overestimate the correct age of the sample. Under normal
circumstances, leaching would start at the surface and be accompanied by lower U-concentrations. While some of the proles show
signicantly lower U-concentrations at or close to the surface, these

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A.S. Mijares et al. / Journal of Human Evolution 59 (2010) 123e132

U-series age estimate (ka)

100
80
60
40
20
0
0

20

40
60
U concentration (ppm)

80

100

Figure 7. Apparent U-series age versus U-concentration (error bars omitted for
clarity).

sections do not show any signicantly older apparent U-series age


estimates. The only region where leaching could perhaps be identied is in the centre of the fourth track between cycles 2190 and
2310 (lower diagram, Fig. 6). The U-concentrations drop by 50%,
while the average apparent U-series results are around 15% older
than the surrounding data without being statistically signicantly
different.
If the bone was younger than 40 ka, the minimum average
amount of uranium leached from it must have been at least 35%,
and the leaching must have happened relatively recently. On top of
this, it must have affected the whole bone in the same manner, i.e.,
parts with high U-concentrations lost 35% of their total uranium,
and the same would have to be lost from parts with low
U-concentrations. Any such process must be regarded as purely
hypothetical, and the above discussion reinforces the need for
developing combined mass spectrometric Pa/UeTh/U analysis of
bones to address questions of U-mobilization (Grn et al., in press).
To summarize, it cannot be excluded that some leaching may
have taken place in the past, causing an increase in the average
apparent U-series results by some unknown amount. However, we
do not nd any clear evidence for leaching, nor could it have been
on a scale sufcient to reduce the minimum age of MT3 to 40 ka or
younger. On balance, we regard the U-age estimate of 66.7  1 ka,
derived from the weighted mean of all data points, as a minimum
age estimate for the MT3. This agrees with the ESR age estimate of
Callao 1, although this may be purely coincidental due to the nature
of the deposit.
Taxonomic identication of the Callao MT3
The general morphology, shape and size of the Callao right MT3
indicate the bone belongs to a primate. For taxonomic identication, morphological comparisons were rst undertaken with
primates of medium to large body size presently living in Island
Southeast Asia (Fig. 8). Even though Macaca and Homo are the only
two extant genera of catarrhines in the Philippines (Fooden, 1991;
Brandon-Jones et al., 2004; Harrison et al., 2006), representatives
of the genera Pongo and Hylobates are added to this comparison.
Indeed, their present geographic extension reaches the neighboring
island of Borneo, and the morphology of their metatarsals is
potentially close to the morphology of the Callao specimen. For
comparison from a palaeontological perspective, the left MT3 of
Homo habilis, OH8, is included in the comparative sample. This
specimen (whose 3D model is mirrored in Fig. 8 for easiest direct
comparison) is part of one of the oldest groups of foot bones
attributed to the genus Homo, and related to a partly or fully bipedal
locomotion (Day and Napier, 1964; Archibald et al., 1972; Kidd et al.,
1996; Berillon, 2000; Susman, 2008; see Wood, 1974; and Gebo and
Schwartz, 2006 for a different taxonomic attribution of OH8).

Several diagnostic features of the Callao metatarsal can be


underlined from this comparison (Fig. 8). In the comparative
sample, the global size of the Callao metatarsal is medium: it is
smaller than Pongo, larger than Hylobates and Macaca, and ts
within the genus Homo. In medial view, the dorsal face of the shaft
of the Callao specimen is straight, as seen in the two representatives of the genus Homo, and not convex as in Pongo, Hylobates, or to
a lesser extent Macaca. In dorsal view, the proportions of the Callao
MT3 shaft, the shape of the base, and the oblique orientation of the
proximal articular facet for the lateral cuneiform are close in shape
to those observed in other Homo specimens. In this view, the
medial and lateral borders of the shaft of the Callao MT3 are slightly
and regularly convergent from the proximal portion to the distal
end of the shaft, which is a condition also encountered in OH8 and
Homo sapiens in general. A different morphology is observed in the
other genera: the medial and lateral borders of the shaft appear
sub-parallel in Pongo, and they are slightly divergent anteriorly in
Hylobates and Macaca. The proximal view shows an outline of the
base of the Callao MT3, which is denitely closer to Homo than to
any other primates included in the comparison. Its elongated
triangular outline is completely different from the outline found in
Pongo, Hylobates and Macaca. In those genera, the medial and
lateral borders of the base have very different sizes and shapes,
with articular facets for MT2 and MT4, which are organized and
orientated differently. In this comparison, the base of the Callao
MT3, which is proportionally shorter in the dorso-plantar direction,
tends to be closer in shape to OH8 than to the Philippine Negrito
specimen used for comparison. However, variability in this feature
is rather common in H. sapiens.
Altogether, these features indicate that the Callao MT3 belongs
to the genus Homo. It is especially close to small-bodied Homo, such
as African H. habilis and present-day Philippine Negritos, the latter
being much closer chronologically and geographically. However,
some differences, such as the reduced proportions of the base, were
observed. They will be discussed further in the morphological and
metrical descriptions that follow.
Age at death
During individual growth, the head is the last anatomical part to
fuse on metatarsals, between the ages of 12 and 16 years on average
in anatomically modern H. sapiens (Scheuer and Black, 2000). The
absence of the head could thus indicate a non-adult individual (i.e.,
with the distal head being unfused at the time of death) or, alternatively, a broken, fully developed and fused adult metatarsal. The
distal extremity of the Callao metatarsal has a fracture, which
occurred at the level of the dorsal tubercles and is obliquely oriented
with respect to the long axis of the shaft. The broken distal
extremity is slightly eroded and partly covered by sediment, which
makes it difcult to ascertain the presence or absence of a metaphyseal surface. Such a surface can also not be detected from analysis of CT-scan data through close-up observation of the CT slices at
the distal extremity (Fig. 9). On the other hand, the morphology and
size of the preserved portion of the medial tubercle seem to be
rather large and fully developed. This would better correspond to an
advanced stage of growth, with a distal head that was about to fuse,
or was already fused at the time of death. These observations point
more to an adult or young adult metatarsal, which was broken
distally after burial. However, it is impossible to discard entirely the
possibility that the specimen is from a juvenile or adolescent.
Morphological and metrical features: comparisons with H. sapiens
As seen in the anatomical comparisons, the general morphology
of the Callao metatarsal matches those observed in the genus Homo

A.S. Mijares et al. / Journal of Human Evolution 59 (2010) 123e132

129

Figure 8. Comparison of (A) the Callao MT3 in proximal, dorsal and medial views with right third metatarsals of (B) Homo sapiens, male Negrito; (C) Homo habilis, OH 8 e mirrored
3D model of a cast of the left MT3; (D) Macaca, (E) Hylobates; and (F) Pongo. 3D models obtained by surface scanning with a NextEngine Desktop 3D Scanner.

and appear close to H. sapiens. Some morphological particularities


also diverge to some extent from the general morphology
encountered in anatomically modern H. sapiens (see Fig. 4 for the
morphological features discussed, and Fig. 8 for comparison with
the MT3 of a male Philippine Negrito).
In proximal view, the shape of the base is typically human. It
shows a triangular contour with slightly depressed medial and
lateral sides, the latter being more depressed just under the articular facet for the fourth metatarsal. However, the medial facet for
the second metatarsal appears to be more plantarly oriented than
generally encountered in H. sapiens. In lateral view, the proportions
and especially the dorso-plantar development of the proximal part
are reduced compared with the size and development of the shaft.
The proximal articular facet for the lateral cuneiform shows
a marked dorso-plantar convexity where it is usually at or slightly
concave in anatomically modern H. sapiens. More precisely, the
convexity of the facet in lateral view is mainly due to the orientation of the plantar portion of the base that is clearly oblique to the
main axis of the bone and thus marks an angle with the dorsal
portion of the base. On the lateral side of the base, the articular

facet for the fourth metatarsal is clearly visible; it is well developed


and slightly elongated. A bulging tubercle is also present, distal to
the lateral articular facet. On the medial side, only one articular
facet for the second metatarsal is visible and located on the dorsal
corner of the base. After cleaning the attached sediment, the
plantar portion of the medial side of the base appears to be rather
at, but no delimited facet is visible. It remains possible that
insufcient cleaning prevents us from observing the plantar facet,
but it is not uncommon in the H. sapiens MT3 for the second facet
for MT2 to be reduced in size or even totally absent. In a study of
100 MT3, Singh (1960) found 41 specimens without this plantar
facet.
The shape of the shaft is also unusual, especially when seen in
lateral view. The dorsal face of the shaft is very slightly convex,
almost straight and horizontal, and with a longitudinal ridge
running along the dorsal surface. In H. sapiens, the plantar surface
of the shaft generally tapers gradually from the base to the head.
Accordingly, the plantar surface around the mid-shaft has a slight
longitudinal concavity, with more marked and abrupt concavities
located just anterior to the base and posterior to the distal head.

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A.S. Mijares et al. / Journal of Human Evolution 59 (2010) 123e132

Figure 9. CT-scan views of the Callao MT3. A: midsagittal CT slice; B: 3D model with transparency (lateral view); C: close-up view of a sagittal CT slice of the distal extremity
(the slice is parallel to A but more medially located). Images A and C show the distal extremity of the MT3, which is broken and partially lled-up by sediment. No transverse
metaphyseal surface is visible.

The Callao metatarsal shows a marked concavity just anterior to the


base and then a more regular and pronounced concavity of the
plantar surface of the shaft from the rst third of its length to
the most distally preserved portion. No clear and marked change in
the angle towards the head is visible. Consequently, the crosssection at the mid-shaft is almost circular, attened medio-laterally
and enlarged in the dorso-plantar direction for the distal-most
third of the shaft.
While not formally developed in this paper, general metrical
comparisons with H. sapiens samples were undertaken with data
from the literature (for instance Volkov, 1903e1904; Gabunia et al.,
2000a); and also with data gathered by one of us (Dtroit) from late
Upper Pleistocene and Holocene H. sapiens from Indonesia and the
Philippines (see Dtroit et al., 2004; Dtroit, 2006). This rst
approach conrmed that one of the most striking features of the
Callao specimen is its overall small size. As a result, it was decided
to focus, for comparative morphological purposes (and not taxonomic identications), on a series of third metatarsals of recent
Negritos from the collections of Muse de lHomme (Musum
national dHistoire Naturelle) in Paris (Table 1). Indeed, Negritos are
small-sized and slightly built Philippine H. sapiens, inhabiting
regions of Luzon Island close to where Callao Cave is located. The
collections from Muse de lHomme also have the advantage of
being well documented and repeatedly studied from various
anthropological perspectives (see, for instance, Genet-Varcin, 1951,
who includes sex determinations of the skeletons; and Volkov,
1903e1904 for the rst published study of the foot bones).
The partial length of the Callao bone, taken parallel to the main
axis of the shaft from the proximal tip of the base to the level of the
medial tubercle, overlaps with the range of variation measured for

male and female Negritos. However, the dimensions of the base of


the bone and the section of the shaft are smaller, indicating peculiar
proportions for the Callao metatarsal. At the mid-shaft, the shaft
appears to be considerably smaller in the dorso-plantar direction
than in the Negrito comparative sample. As shown by the reduced
dimensions obtained for the dorso-plantar height and mediolateral breadth of the proximal facet for the lateral cuneiform, the
base is very small. It is the smallest of our sample, conrming the
particular shape and proportions of the bone as seen from lateral
and superior views.
Discussion
For more detailed future comparisons, it is important to take
into account the paucity of Southeast Asian Homo erectus and
Upper Pleistocene H. sapiens postcranial remains. The fossil record
is particularly decient in foot bones, with none recorded so far for
Southeast Asian H. erectus (Widianto, 1993; Antn, 2003). For
H. sapiens, several rather complete late Upper Pleistocene and
Holocene human skeletons have been recovered from Indonesia
(Jacob, 1967; Dtroit, 2002, 2006; Simanjuntak, 2002; Smah et al.,
2004), and a few fragmentary postcranial remains come from the
Philippines, including foot bones (Dtroit et al., 2004). For those
specimens we are able to observe and measure, the dimensions
largely exceed the dimensions of the Callao metatarsal, and the
proportions of the base are never reduced to the extent seen in the
Callao metatarsal.
From broader taxonomic, chronological and geographic
perspectives, metatarsal bones appear to have been relatively
stable in terms of overall morphology and size since the rst African

A.S. Mijares et al. / Journal of Human Evolution 59 (2010) 123e132

131

Table 1
Comparative measurements of the Callao metatarsal with Negrito samples
Total length

Partial length
(from the base to
the medial tubercle)

Shaft
Dorso-plantar
diameter

52.58

Negritos males (n 11)


Mean
64.69
Std error
0.862
Min.
59.5
Max.
68.3
Median
65.2
Negritos females (n 12)
Mean
59.21
Std error
0.554
Min.
56.5
Max.
62.7
Median
58.55

Callao

60.99a

Proximal facet for lateral


cuneiform

Facet 1 (dorsal)
for metatarsal 2

Facet for
metatarsal 4

Medio-lateral
diameter

Dorso-plantar
height

Length

Height

Length

6.58

5.99

11.28

8.69

5.76

3.82

6.46

4.92

55.66
1.033
49.4
60.5
56.8

8.51
0.107
8
9.1
8.5

7.32
0.268
6.5
9.6
7.1

18.79
0.427
16
21.1
18.8

11.72
0.245
10.2
13.3
11.7

6.46
0.418
4.5
8.9
6.3

4.74
0.269
3.2
6.3
4.7

8.02
0.277
7
10.2
8

7.73
0.323
6
9
7.7

51.33
0.357
49.2
53.5
51.15

7.33
0.091
6.9
7.8
7.35

6.42
0.179
5.1
7.3
6.45

16.87
0.209
15.5
17.9
16.95

10.88
0.211
9.5
12.3
10.8

6.54
0.198
5
7.4
6.8

4.95
0.131
4.3
5.9
4.8

8.06
0.353
6.2
10.5
7.9

7.98
0.329
5.7
9.9
8

Medio-lateral
breadth

Height

a
Estimated from the regression of total length against partial length for the pooled-sex sample of 23 adult Negrito metatarsals III (r 0.93, p < 0.01;
Ltot 1.022Lpart 7.2519).

representatives of the genus Homo (i.e., H. habilis and Homo


rudolfensis). Differences in function have been widely discussed for
the morphology and anatomical position of MT1, as well as the
implication of metatarsals in the development of longitudinal and
transverse plantar arches (see for instance Aiello and Dean, 1990;
Berillon, 2000; Zipfel et al., 2009). As far as the third metatarsal is
concerned, the particular form (or relative proportions) of the base
and shaft of the Callao bone has apparently not been described so
far from other early fossil specimens of the genus Homo from
Eastern Africa, Southern Europe, or the Caucasus. This observation
is conrmed by direct comparison with the H. habilis left MT3 of
OH8 (Fig. 8, and see Day and Napier, 1964; Kidd et al., 1996; Berillon,
2000), and from comparisons with published data from Atapuerca
Gran Dolina (Lorenzo et al., 1999) and Dmanisi (Gabunia et al.,
2000a,b; Lordkipanidze et al., 2007).
Interestingly, the remains discovered at Liang Bua in Flores,
representing several individuals attributed to the species H. oresiensis, include foot bones, and especially the complete series of
right and left metatarsals of the individual LB1 (Brown et al., 2004;
Morwood et al., 2005; Jungers et al., 2009a). The length of the left
MT3 (LB1/23), at 60.4 mm (Jungers et al., 2009b), is near the estimated length for the Callao specimen (60.99 mm; see Table 1).
Although few details are available regarding the exact morphology
and shape of this particular bone, it certainly calls for interesting
future comparisons. It is very difcult to determine accurately
a taxonomic attribution from a single bone and, with the exception
of the Liang Bua remains, the MT3 recovered from Callao Cave on
Luzon Island is the only Upper Pleistocene foot bone attributed to
the genus Homo discovered beyond Wallaces Line in Island
Southeast Asia. Additionally, the preliminary description of the
Callao MT3 clearly indicates a reduced general size only known in
this region in H. oresiensis and in small-bodied H. sapiens, such as
present-day Philippine Negritos.

Provisionally attributed to a rather small-bodied H. sapiens, the


Callao MT3 documents the presence of a hominin species on the
island of Luzon as early as 67 ka, and is testimony to a capability to
colonize new territories across open sea gaps. The Philippine
specimen also indicates that Flores was not the only island in
Wallacea to be occupied by hominins more than 50,000 years ago.
The discoveries at Callao Cave also raise some other important
questions about the cultural behavior of these early colonizers of
the Philippine archipelago. For example, even though there is
evidence of butchery in the animal bone assemblage, not a single
stone implement was recovered, suggesting perhaps the use of an
alternative technology. To answer such questions, more archaeological research in the caves of karstic limestone formations of the
Philippines is required.
Acknowledgement
We are grateful for the support of the National Museum of the
Philippines, the Cagayan Provincial Government, and the Protected
Area Management Board-Peablanca. Roberto Macchiarelli and
Arnaud Mazurier are acknowledged for the CT-scan of the bone. We
are also grateful to Eduardo Bersamira and Myra Lara for the
illustrations, and Archie Tiauzon as a member of the excavation
team. Funding for this research came from an Australian Research
Council Discovery Grant to Peter Bellwood and a University of the
Philippines Research Grant to Armand Mijares. Florent Dtroit
received support from Asia-Link HOPsea and Erasmus Mundus
Action 3 exchange programs. We are grateful for the support of
Philippe Mennecier and Aurlie Fort for the access to the collections
at the Muse de lHomme. The Ofce of the Vice Chancellor for
Research and Development, University of the Philippines funded
Philip Piper. The dating aspects of this study were supported by
Australian Research Council Grant DP0664144 (Grn et al.) Microanalysis of human fossils: new insights into age, diet and migration.

Conclusions
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A comprehensive analysis of size and shape characteristics,
including comparisons with larger samples of H. sapiens and fossil
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