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BIOLOGY

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RIES NO. 42

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e Contribution of Archaeology
to the Zoogeography of Borneo,
with the First Record of a Wild Canid
of Early Holocene Age

Earl of Cranbrook

A Contribution in Celebration

of the Distinguished Scholarship of Robert F. Inger

on the Occasion of His Sixty-Fifth Birthday

HALL

March 31, 1988
Publication 1385

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

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Croat, T. B. 1978. Flora of Barro Colorado Island. Stanford University Press, Stanford, Calif, 943 pp.
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FIELDIANA

Zoology

NEW SERIES, NO. 42

The Contribution of Archaeology
to the Zoogeography of Borneo,
with the First Record of a Wild Canid
of Early Holocene Age

Earl of Cranbrook

Great Glemham House
Saxmundham
Suffolk. IP 17 1LP
England

A Contribution in Celebration

of the Distinguished Scholarship of Robert F. Inger

on the Occasion of His Sixty-Fifth Birthday

Accepted for publication March 31, 1986
March 31, 1988
Publication 1385

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

© 1988 Field Museum of Natural History

ISSN 0015-0754

PRINTED IN THE UNITED STATES OF AMERICA

Table of Contents

List of Tables

Abstract 1

Introduction 1

Canid Remains from Agop Sarapad, Ma-

dai Caves, Sabah 2

The Specimens 3

Discussion 4

Identification of the Specimens 4

Post-Pleistocene Extinctions in Borneo ... 5

Acknowledgments 6

Literature Cited 6

Measurements of the left calcaneum of
selected specimens of canid species

List of Illustrations

The dhole calcaneum and canine tooth
from Agop Sarapad, Madai caves, Sa-
bah 2

The calcaneum positioned in calipers to
measure maximum length 4

in

The Contribution of Archaeology
to the Zoogeography of Borneo,
with the First Record of a Wild Canid
of Early Holocene Age

Abstract

The extant fauna of Borneo lacks several large
mammal species that are widespread elsewhere in
the Sunda region. Archaeological research in Bor-
neo has already found remains of three of these in
late Upper Pleistocene or Holocene contexts: the
tiger, Panthera tigris; Malay tapir, Tapirus indicus;
and Javan rhinoceros, Rhinoceros sondaicus. Evi-
dence of a fourth, the dhole, Cuon alpinus, is now
reported from a midden dated about 10,000 b.p.
in Agop Sarapad, Madai caves, Sabah. Extinction
of these species evidently occurred within the last
few thousand years and is attributed to failure to
adapt to the environmental consequences of post-
glacial climatic changes.

Introduction

In his monograph on the amphibians of Borneo,
Robert Inger anticipated that new species would
be discovered but that these additions would not
materially affect conclusions on the geographical
relations of the fauna (Inger, 1966, p. 357). Both
predictions have since come true. Inger himself
has subsequently described new Bornean taxa (In-
ger & Frogner, 1979; Inger & Gritis, 1983), and
Dring (1983a,b) has added others. Yet the overall
picture of amphibian zoogeography in the region
remains fundamentally unaltered. The fauna un-
doubtedly originated from continental Asia and
shows little affinity with Celebes, only a short dis-
tance to the west, or with the Philippine Islands.
There has, however, been a significant local ra-
diation at lower taxonomic levels, recognizable in

an assemblage of species confined to the Sundaic
region; that is, the islands of the Sunda shelf and
the Malay Peninsula south of about 10°N latitude.
Species endemic to Borneo (comprising about 40%
of the island's amphibian fauna) represent a spe-
cial group among the Sundaic assemblage. Of the
nonendemics, many species (about 80%) also oc-
cur in Sumatra, somewhat fewer in Peninsular Ma-
laysia (formerly Malaya), and only about 45% in
Java.

Similar patterns characterize the zoogeography
of the other terrestrial vertebrate classes. Thus,
among the mammals, the fauna has clearly orig-
inated from continental Southeast Asia, modified
by a significant radiation within the Sunda region.
Endemism is high in Borneo, with 37 (29%) of 1 29
species of land mammals (i.e., excluding bats and
marine mammals) currently being recognized as
endemics. Of the 92 nonendemic species in Bor-
neo, 87 (95%) also occur in Sumatra, 80 (87%) in
Peninsular Malaysia, and only 44 (49%) in Java
(data from Medway, 1977a, and Zon, 1979).

Within the Sunda region, anomalous distribu-
tions can be recognized, in several cases involving
large mammal species which might be expected
most easily to cross geographic barriers. In con-
temporary Borneo, notable absentees are tiger,
Panthera tigris, at present in Peninsular Malaysia
(pm), Sumatra (s), and Java (j); leopard, Panthera
pardus (pm, ?s, j); wild dog or dhole, Cuon alpinus
(pm, s, j); Eurasian wild pig, Sus scrofa (pm, s);
Javan rhinoceros, Rhinoceros sondaicus (pm, s, j);
Malay tapir, Tapirus indicus (pm, s); and serow,
Capricornis sumatraensis (pm, s). Among smaller
mammals, only one giant squirrel of the genus
Ratufa occurs (i.e., R. affinis), the widespread R.
bicolor (pm, s, j) being absent. Also lacking are the

CRANBROOK: ZOOGEOGRAPHY OF BORNEO

bamboo rat, Rhizomys sumatrensis (pm, s), and
the brush-tailed porcupine, Atherurus macrourus
(pm, s). The banteng, Bos sondaicus, the only wild
cattle in Borneo, is also native in Java, but is ab-
sent from Sumatra and Peninsular Malaysia. Like
the ferret badger, Melogale orientalis (j), B. son-
daicus occurs in parts of mainland continental
Southeast Asia, but only in Java and Borneo on
the Sunda shelf. These two distributions, coupled
with the occurrence in Borneo of a species of long-
nosed ground squirrel, Dremomys, and a smooth-
tailed treeshrew, Dendrogale (genera found else-
where only on continental Southeast Asia), led
Chasen (1940, pp. xi-xv) to propose a limited col-
onization of the Sunda region by "eastern drift"
from the Indochinese region. He also envisaged a
"western drift from the continent by way of Su-
matra" to explain the absence of the Eurasian wild
pig, tiger, and dhole from Borneo.

Archaeological evidence shows that neither pos-
tulate is in fact required. It has long been known
that the Pleistocene mammal fauna of Java in-
cluded species now extinct on that island but sur-
viving elsewhere in the Sunda region; for example,
siamang, Hylobates syndactylus; orangutan, Pongo
pygmaeus; Malay bear, Helarctos malayanus; el-
ephant, Elephas maximus; bearded pig, Sus bar-
batus; tapir; and bamboo rat. In Sumatra, banteng
remains have been found in cave deposits attrib-
uted either to a Middle Pleistocene interglacial or
to the early Holocene (data reviewed by Hooijer,
1975, and Vos, 1983). More recently, excavations
by the Sarawak Museum in the 1950-1 960s and
the Sabah Museum in the early 1980s have re-
vealed the former presence in Borneo of four of
the large mammals missing from the modern fau-
na: (1) the tiger, represented at Niah cave, Sarawak
(see Harrisson, 1972, and bibliography therein),
by a deciduous canine from Neolithic levels
(Hooijer, 1963); (2) the tapir, represented at Niah
by a dozen pieces ranging in date from late Upper
Pleistocene to about 8000 b.p. (Medway, 1960);
(3) the Javan rhinoceros represented by a decid-
uous molar and a fragment of ulna in a midden
dated at about 10,000 b.p. (designated "MAD 2"
by the excavator) in the Agop Sarapad mouth of
Madai caves, Sabah (Bellwood, 1984), and (iden-
tified with lessened confidence) by a left ectocu-
neiform and a fragmentary lateral proximal pha-
lanx from depths at Niah attributable to an early
Holocene age (see Cranbrook, 1986, for details of
these rhinoceros finds); and (4) the dhole, docu-
mented below. The problem in these cases is no
longer to elucidate routes or barriers to the colo-

nization of Borneo, but rather to explain local ex-
tinctions within the past few thousands of years.

Canid Remains from Agop Sarapad,
Madai Caves, Sabah

The excavations were carried out in 1980 in the
caves at Madai near Kunak, Sabah, 118°08'E,
4°44'N, by staff of the Sabah Museum in associ-
ation with Dr. Peter Bellwood (Bellwood, 1984,
fig. 1; Bellwood, unpubl. data). Animal remains,
including vertebrate teeth and bones, were sepa-
rately bagged from each layer and later examined
by myself in the museum premises at Kota Kin-
abalu. Notes made at the time have been worked
into a general report on animal remains, which
has been placed on file in the museum. It is planned
to publish an edited version as part of the exca-
vator's final report (Bellwood, unpubl. data). Spec-
imens appearing to be of particular interest (and
therefore brought to England for further study)
included the two described below. Both derive from
the Agop Sarapad mouth (MAD 2), Madai caves,
square HI, layer 2, at 5-10 cm. This provenance
is firmly within the MAD 2 riverine shell midden,
14C-dated to about 10,000 b.p. (Bellwood, pers.
comm.). The specimens are illustrated in Figure
1 . Both have been returned to the Sabah Museum,
Kota Kinabalu, Malaysia, for safekeeping. A cast

Fig. 1 . The dhole calcaneum and canine tooth from
Agop Sarapad, Madai caves, Sabah. Scale is in centi-
meters.

FIELDIANA: ZOOLOGY

Table 1 . Measurements of the left calcaneum of selected specimens of canid species.

Measurementst (mm)

Minimum

BM(NH)

Median

Maximum

Maximum Maximum

breadth

Item*

Sex

reg. no.

length

length

depth

breadth

(tuber)

Archaeological specimen

Agop Sarapad. Madai

M42838t

43

45.5

18.3

17.6

7.2

Malaysian pariah-type dog

("Manggis")

F

71.753

35.5

36.6

15.2

14.3

6.0

Cuon alpinus javanicus

M

1888.2.5.22

41.8

43.9

18.6

16.8

7.4

Cuon alpinus dukhunensis

F

1936.4.8.1

43.7

45.2

19.0

18.8

8.5

Canis familiaris dingo

9

1954.11.15.1

40.8

43.5

17.5

16.4

7.3

Canis familiaris dingo

F

1868.4.14.1

46.2

47.8

19.2

17.3

6.9

Canis lupus

7

1937.2.10.2

49.5

52.2

21.2

22.0

8.6

Canis aureus anthus

9

816c

38

40.8

14.4

15.5

5.6

* All examples are full-grown adults in the collection of the British Museum (Natural History).

t Measurements were taken as follows: median length — from the tendinial groove on the superior face of the tuber
calcanci to the inferior facet of the body; maximum length— the greatest length measured by calipers positioned as
shown in Figure 2; maximum depth— the greatest distance between the anterior and posterior surfaces measured at
the base of the tuber; maximum breadth— the breadth from the median edge of the sustentaculum tali to a point
opposite it on the lateral surface on the body of the calcaneum; minimum breadth (tuber)— the mediolateral breadth
of the tuber calcanei at its narrowest point.

% A cast. The original has been returned to the Sabah Museum.

of the calcaneum has been made by the British
Museum (Natural History), where it is held in the
Palaeontology Department under the registration
number M 42838.

The Specimens

Right Lower Canine— Damaged, now con-
sisting only of the root and dentine core of the
crown. A very small area of enamel remains on
the outer (labial) face at the base of the crown,
shiny dark brown in color. This small piece of
enamel is adequate to provide an orientation point
from which the root can be measured, but is of no
value for the specific identification of the tooth.

Measurements are as follows:

Root, length 23.3 mm

Root, maximum breadth in anteroposterior

(= mesiodistal) plane 9.7 mm

Root, maximum breadth in lateral plane

6.5 mm

Left Calcaneum— Complete and undamaged.
This bone is colored dark brown. It is indisputably
attributable to a member of the dog family, Can-
idae. Among likely species of Asian wild dogs,
comparative measurements (table 1 ) show that this
calcaneum is slightly larger in principal dimen-
sions than its homologue in the skeleton of an
adult male dhole of the Javan subspecies, Cuon

alpinus javanicus, but smaller than that of an adult
female of the Indian subspecies, C. a. dukhunensis.
It is distinctly smaller than wolf, Canis lupus, and
larger than jackal, Canis aureus (a member of the
African race, C. aureus anthus, being the only
available adult example). Neither is therefore in-
dicated by the measurements. The discovery of
either of these two Canis species in Borneo would
represent a large extension of known range, his-
toric or prehistoric. Both are thus best excluded
from consideration.

There remains the possibility that a type of do-
mestic dog Canis "familiaris" is represented.
Available for comparison is the skeleton of a fe-
male of the primitive Malaysian pariah-type breed
of domestic dog, an individual described and pic-
tured by Medway (1977b, plate 3, 'Manggis'), now
deposited at the British Museum (Natural Histo-
ry); this is markedly smaller. Examples of Austra-
lian dingo in the bm(nh) collections show wide
variation in size; measurements of the calcanea of
two adults bracket the archaeological specimen.

Examination of a wider selection of calcanea of
dholes and dingos does not reveal any nonmetric
character that serves to distinguish the species.
Features investigated included the relative dimen-
sions of the tuber calcanei, the shape and angle of
the sustentaculum tali, the angle of the ventral face
of the process at the anterior base of the tuber,
and the size and shape of the small facet at the
mesioventral comer of the anterior face of the body

CRANBROOK: ZOOGEOGRAPHY OF BORNEO

«

Fig. 2. The calcaneum positioned in calipers to measure maximum length (table 1).

of the calcaneum (terminology following Hughes
& Dransfield, 1953). In no case was there any
species-specific difference discernible in the sam-
ples available. I conclude that, on morphology
alone, the specimen could be attributed to a dhole
or to a primitive, dingo-like form of domestic dog.
Arguments for a more positive identification must
be based on nonmorphometric grounds.

Discussion

Identification of the Specimens

On the one hand, the dog was among the first
of the mammals to be domesticated. In Europe
and the Near East, remains attributed to dogs,
rather than wolves, appear in association with Epi-
paleolithic or Mesolithic cultures of early postgla-
cial age. Dates suggested for the pioneer successes
in domestication range from 12,000 b.p. (Davis &
Valla, 1978) to 14,000 b.p. (Nobis, 1979).

On the other hand, no such early dates have
been established in Southeast Asia. In Borneo, re-
mains of undoubtedly domesticated dogs have been
found in archaeological contexts in other caves,
but in none have cultural associations indicated

an age earlier than Neolithic, at the oldest (Med-
way, 1977b). In Thailand prehistoric sites have
produced domestic dog bones dating back to 3500
b.c. (Higham et al., 1980), a period considerably
less ancient than the Agop Sarapad midden.

Relevant evidence is the lack of other signs of
the presence of domestic dog at these sites. While
examining the bone from Agop Sarapad (and also
the much larger quantity from deep levels at Niah),
I have looked for the distinctive marks of gnawing
by carnivores and found none. These middens
originated as the food remains of human cave vis-
itors. If contemporary man had been accompanied
by domestic dogs, at least some of the bones must
have been gnawed. I conclude that the most plau-
sible identification for the two archaeological spec-
imens is a wild dog, which itself was perhaps tl{e
quarry of man. On the grounds of size and zoo-
geography, the dhole is indicated.

Intriguingly, as for tiger (Gersi, 1975) and tapir
(Medway, 1977a), anecdotal accounts exist of the
presence of the dhole in Borneo in historic times.
Hose (1893, p. 26) included Borneo in the range
of the species (under the name Cyon rutilans),
commenting: "This wild dog must be very rare in
Borneo. I have constantly heard native accounts
of it, but I have never seen a specimen." Later,
recounting a trip to Batu Bukit Song, Sarawak,

FIELDIANA: ZOOLOGY

Hose (1929, p. 144) reported sighting a "pair of
wild dogs . . . they had been eating the remains of
a young wild pig, but before we had time to get
more than a glimpse of them they were away." At
no time in the history of zoological collection has
a specimen been taken. Hose's record was not ad-
mitted by subsequent authors, including E. Banks
and F. N. Chasen (see Chasen, 1940, p. 93), both
of whom had much field experience in the region.

Post-Pleistocene Extinctions in Borneo

The complete list of 58 mammal species iden-
tified in the excavations at Niah, including do-
mestic dog and goat but excluding the Javan rhi-
noceros (at that time unrecognized), has been
published elsewhere (Medway, 1979, table 2). The
total from Madai is smaller (Cranbrook, unpubl.
ms. deposited at Sabah Museum, Kota Kinabalu).
Both lists serve to confirm that the mammal fauna
at the threshold of the Holocene era was very sim-
ilar to that of present-day Borneo.

The deepest levels at Niah yielded remains of
one totally vanished mammal, the giant pangolin,
Manis palaeojavanica (Hooijer, 1 960a), but this
apparently did not survive into the Holocene era.
The early Holocene fauna of Borneo presumably
included the mammals already mentioned (tiger,
dhole, tapir, and Javan rhinoceros) which have
since become locally extinct. Over the same pe-
riod, other species, which persist in modern Bor-
neo, have undergone metrical changes. Thus, there
has been a decline in tooth size among orangutans
(Hooijer, 1960b) and monkeys of the genera Pres-
bytis (including subgenus Trachypithecus) and
Macaca (Hooijer, 1962). Among two species of
giant rats, Rattus sabanus and R. muelleri, there
have been changes both in the size and the relative
dimensions of the teeth (Medway, 1964). Larger
body size among the prehistoric populations is
indicated by the dimensions of limb and foot bones
of Sumatran rhinoceroses, Dicerorhinus suma-
trensis (Cranbrook, 1 986), and barking deer, Mun-
tiacus muntjak (Medway, 1959).

The same archaeological studies show that man
was present throughout the period concerned. Yet,
as I have argued elsewhere (Medway, 1979), there
are no grounds to suggest that direct or indirect
impacts of human activity contributed to evolu-
tionary changes or to the extinction of any species
before the advent of the shotgun. Explanations
must be sought in natural ecological processes.

Advances in palaeogeographical research have

improved our understanding of events in the re-
gion during the Cenozoic and subsequently. The
antiquity of the Makassar Strait is confirmed, al-
though it is now recognized that Celebes has a
complex tectonic origin (Audley-Charles, 1981).
During the Pleistocene there were large extensions
in the exposure of subaerial land of the Sunda shelf
associated with glacial episodes, as summarized
by Inger (1966) from earlier sources. It is also
known that these were accompanied by climatic
changes sufficient, for instance, to cap Mt. Kina-
balu with permanent ice (Koopmans & StaufTer,
1967). It seems that present-day conditions in the
Sundaic region are atypical of the Quaternary as
a whole, with sea level and the upper forest limit
on mountains exceptionally high, average tem-
peratures close to maximum Quaternary values,
and the climate at or near its wettest (Whitmore,
1 98 1 ). A change from cooler, drier, and more sea-
sonal conditions apparently took place rather rap-
idly in the first millenia of the Holocene, accom-
panied by rising sea levels which reached a
maximum of at least 6 m above present shorelines
about 5000 b.p. (Haile, 1971).

The cooler and more seasonal climate of Pleis-
tocene glacial periods, including the last, which
immediately preceded the Holocene intermission,
is presumed to have favored the development of
a mosaic of forest broken by open gaps and glades.
Under such conditions, the forest-edge facies must
have been extensive, with abundant browse ac-
cessible to ground-dwelling ungulates. Following
the change to a warmer and wetter postglacial cli-
mate, the open spaces would have been invaded
by closed forest. Mature \ropical evergreen rain
forest supports a sparse ground and shrub storey
vegetation. In the deep shade, plant life is restrict-
ed. The main zone of productivity is in the upper
canopy, high out of reach of ground-dwelling her-
bivores. These changes would therefore have de-
graded the quality of the environment for all non-
scansorial phytophagous mammals of the ground
or shrub storeys, and for large-bodied browsing or
grazing perissodactyls in particular. Indirect con-
sequences would have impinged upon nonscan-
sorial predators of these mammals, again espe-
cially the large-bodied carnivores. The affected
populations would thus have experienced ecolog-
ical pressures simultaneously with the final sev-
erance of land connections across the Sunda shelf
and hence isolation from sources of new genetic
inflow.

Nevertheless, Bornean populations of other
mammal species have continued from the early

CRANBROOK: ZOOGEOGRAPHY OF BORNEO

postglacial to the present. Among these, the beard-
ed pig, Sus barbatus, as far as archaeological ma-
terial can show, has undergone no morphological
change (Medway, 1978). At the same time, the
Sumatran rhinoceros and barking-deer have re-
sponded to the changing environment by reduc-
tion in individual body size, a modification rec-
ognized as adaptative to restricted resources. The
metrical changes in the teeth of other mammals,
as noted above, involving absolute and/or relative
dimensions, may also reflect alterations in body
size.

The archaeological evidence is inadequate to
demonstrate changes in abundance. Populations
affected by postglacial alterations in the environ-
ment were likely to have declined in response to
the diminution of accessible resources. Large
mammals might already have been comparatively
rare and hence vulnerable to local extinction, even
on so big an island as Borneo. Reduced population
density among the primary consumers has inevi-
table repercussions on members of the community
at higher trophic levels. It is therefore not unex-
pected that the mammal species to become extinct
in Borneo during the postglacial period included
these four obligatory ground-dwellers: two large
browsing ungulates (Javan rhinoceros and tapir)
and two large predatory carnivores (tiger and
dhole).

I would be venturing too far from my own ground
if I were to speculate on the likely effects of these
environmental changes on amphibian faunas of
the terminal Upper Pleistocene of Borneo. This
train of thought I hereby offer to Bob Inger, with
respect and affection, leaving him to pursue it as
far as may be profitable in the light of his own
studies of the existing fauna.

Acknowledgments

I am grateful to Dr. Peter Bellwood for supplying
radiometric dates, to Dr. Juliet Jewell for provid-
ing facilities for this study at the British Museum
(Natural History), and to both, with Dr. D. A.
Hooijer, for kindly reading and commenting on
this text in progressive stages of drafting.

Literature Cited

Audley-Charles, M. G. 1981. Geological history of
the region of Wallace's line, chap. 4. In Whitmore, T.

C, ed., Wallace's Line and Plate Tectonics. Clarendon
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Bellwood, P. 1984. Archaeological research in the
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CRANBROOK: ZOOGEOGRAPHY OF BORNEO

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