Megafaunal extinctions and their consequences in the tropical Indo-Pacific

Abstract

The global Quaternary Megafauna Extinction (QME) event eliminated two-thirds of all mammal genera and half (c. 178) of all species of body mass >44 kg, with most well-dated extinctions occurring between c. 50,000 and 3000 years ago (Barnosky 2008). The QME differed from other extinction events in the fossil record by the absence of replacements, the fact that similar episodes occurred in widely separated sites at different times, and the fact that the timing can often be loosely linked with the arrival of modern humans. The causes and consequences of this event have been debated at length in the literature for Australia, Madagascar, northern Eurasia, and both North and South America, with Africa often used as a ‘control’ in comparisons (Koch and Barnosky 2006; Barnosky 2008; Johnson 2009), but tropical Asia, Wallacea (the Indonesian Islands separated by deep water from the Asian and Australian continental shelves) and New Guinea have generally been omitted from these debates. This partly reflects uncertainties in the fossil and archaeological records of the region – in particular, about the timing of large vertebrate extinctions in relation to the arrival first of early Homo and then of modern humans (Bird et al. 2004; Louys et al. 2007; Corlett 2009a). However, the Asian mainland, all large islands and many smaller ones supported more large vertebrate species and much more large vertebrate biomass in the Pleistocene than they do now (Long et al. 2002; Louys et al. 2007; Louys 2008) and an ongoing extinction episode threatens all the survivors (Corlett 2007). Here, I ask what has been lost, when and why, and what the ecological consequences have been.

Full text

Megafaunal extinctions and their consequences in the tropical Indo-Pacic 117
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8
Megafaunal extinctions and their
consequences in the tropical Indo-Pacic
Richard T. Corlett
Department of Biological Sciences, National University of Singapore, Singapore
corlett@nus.edu.sg
Introduction
e global Quaternary Megafauna Extinction (QME) event eliminated two-thirds of all
mammal genera and half (c. 178) of all species of body mass >44 kg, with most well-dated
extinctions occurring between c. 50,000 and 3000 years ago (Barnosky 2008). e QME
differed from other extinction events in the fossil record by the absence of replacements, the
fact that similar episodes occurred in widely separated sites at different times, and the fact
that the timing can often be loosely linked with the arrival of modern humans. e causes
and consequences of this event have been debated at length in the literature for Australia,
Madagascar, northern Eurasia, and both North and South America, with Africa often used
as a ‘control’ in comparisons (Koch and Barnosky 2006; Barnosky 2008; Johnson 2009),
but tropical Asia, Wallacea (the Indonesian Islands separated by deep water from the Asian
and Australian continental shelves) and New Guinea have generally been omitted from these
debates. is partly reflects uncertainties in the fossil and archaeological records of the region
– in particular, about the timing of large vertebrate extinctions in relation to the arrival first of
early Homo and then of modern humans (Bird et al. 2004; Louys et al. 2007; Corlett 2009a).
However, the Asian mainland, all large islands and many smaller ones supported more large
vertebrate species and much more large vertebrate biomass in the Pleistocene than they do
now (Long et al. 2002; Louys et al. 2007; Louys 2008) and an ongoing extinction episode
threatens all the survivors (Corlett 2007). Here, I ask what has been lost, when and why, and
what the ecological consequences have been.
Who was where when?
e earliest human remains in the region are assigned to Homo erectus. Most skeletal
remains are from Java, 1.0-1.8 million years ago, but the earliest securely dated stone tools
are from the island of Flores (Morwood et al. 1998) and South China (Hou et al. 2000),
only 800,000 years ago. Most of the faunal remains associated with H. erectus finds suggest

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Altered Ecologies: Fire, climate and human inuence on terrestrial landscapes118
open woodland or savanna and it is not clear whether the species could live in closed forest.
Other unknowns are its ability to manipulate fire and to make voluntary sea crossings,
such as those needed to reach Flores. Later Asian populations of H. erectus appear to have
diverged increasingly from their African ancestors, and mainland East Asia was occupied
between about 500,000 and 70,000 years ago by hominids who may have represented
evolutionary developments from local H. erectus populations and/or archaic forms of H.
sapiens and/or additional Homo species (Bacon et al. 2006; Louys et al. 2007). A remarkable
dwarf hominin, H. floresiensis, inhabited a limestone cave on Flores from 95 ka to 16.6 ka
(Moore et al. 2009).
ere is no clear archaeological evidence for anatomically and culturally modern humans
in Eurasia before about 45,000-50,000 years ago. is fits with the growing amount of
archaeological, genetic, pollen and charcoal evidence in support of a single dispersal event
that brought modern humans from Africa, where they had emerged 150,000-200,000
years ago, along the coastlines of South and Southeast Asia, to New Guinea and Australia,
within a period of a few thousand years (Mellars 2006; Pope and Terrell 2008). is ‘coastal
express train’ model of modern human dispersal assumes that the pioneer populations
lived initially on coastal resources, moving on as those resources became depleted, and only
later moved inland.
When was what where?
ere is no clear functional justification for a 44 kg cut-off in the definition of megafauna
(Martin and Klein 1984; Hansen and Galetti 2009), but this limit has been widely used in the
literature and is followed here for convenience. Owen-Smith’s 1000 kg minimum has much
stronger functional justification (Owen-Smith 1988), but includes few Pleistocene taxa in the
region between tropical China and New Guinea covered here. Holocene extinctions of smaller
(<44 kg) mammals (only island rats and bats in this region) are covered by Turvey (2009b).
e only known terrestrial megafaunal or near-megafaunal species on the tropical islands
east of New Guinea were the land turtles, terrestrial crocodiles and giant flightless birds that
persisted into the Holocene on New Caledonia and Fiji (Turvey 2009a).
It would be interesting to examine the large vertebrate fauna of the region over the
full period of hominin occupation, but the problems of interpretation of the fossil record
increase rapidly as one goes back in time, so this review is limited to the past million years,
with the main focus on the late Pleistocene, from c. 130 ka, and the Holocene (Table 1).
is summary builds on those by Long et al. (2002), Louys et al. (2007) and Louys (2008),
updated where necessary. I have checked the primary sources wherever possible and taken
a critical stance when examining the Asian literature, excluding some doubtful records.
Scientific names of all taxa are given in Table 1. is table underestimates the regional
loss of megafaunal species because late Pleistocene fossils have usually been assigned to
the nearest living relative, or to a single extinct taxon, while recent molecular studies have
often divided extant taxa (e.g. orangutans, clouded leopards) into geographically separated
species. It also gives a very misleading impression of the extent of local megafaunal losses,
since many large vertebrates still persist somewhere in the region, despite occupying <10%
of their maximum Holocene ranges and an even smaller percentage of their maximum
Pleistocene ranges.

Megafaunal extinctions and their consequences in the tropical Indo-Pacic 119
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Taxon Maximum range Last known extant Mass (kg) Common name
SQUAMATA
Varanidae Varanus spp. Wallacean islands late Pleistocene ?? monitor lizards
Varanus komodoensis Flores, Komodo etc. extant <165 Komodo dragon
V. salvadorii New Guinea extant <50? crocodile
monitor
Pythonidae Morelia spp. New Guinea extant ?? pythons
Python molurus S. China-Sulawesi extant <140 Burmese python
P. reticulatus SE Asia-Wallacea extant <160 reticulated
python
TESTUDINATA
Testudinidae Geochelone spp. Wallacean islands Pleistocene ?? giant tortoises
CASUARIIFORMES
Casuariidae Casuarius casuarius New Guinea extant 30-60 southern
cassowary
C. unappendiculatus New Guinea extant 40-60 northern
cassowary
DIPROTODONTIA
Diprotodontidae Hulitherium New Guinea after 40,000 BP 150
Maokopia New Guinea after 40,000 BP 100
Zygomaturus New Guinea after 40,000 BP 300
Macropodidae Protemnodon spp. New Guinea after 40,000 BP 45-100 kangaroo
PROBOSCIDEA
Elephantidae Elephas spp. Sulawesi, Luzon Pleistocene ?? dwarf elephants
Elephas maximus China to Java extant 2500-
4000
Asian elephant
Palaeoloxodon
naumanni
China, Vietnam late Pleistocene ?? straight-tusked
elephant
Stegodon spp. China to Timor mid P. to Holocene 300-
3000?
stegodons
PRIMATES
Hominidae Pongo spp. China to Java late P. to Holocene >50 orangutans
P. abelii Sumatra extant 40-90 Sumatran
orangutan
P. pygmaeus Borneo extant 30-90 Bornean
orangutan
PHOLIDOTA
Manidae Manis sp. Borneo 40,000 BP 50? giant pangolin
CARNIVORA
Felidae Panthera pardus China to Java extant 30-70 leopard
P. tigris China to Java extant 75-250 tiger
Hyaenidae Crocuta crocuta China to ailand extant (Africa) 45-80 spotted hyena
Pachycrocuta
brevirostris
China to Java late Pleistocene? >100 giant hyena
Ursidae Ailuropoda
melanoleuca
China to ailand extant 70-115 giant panda
Table 1. Terrestrial vertebrates with body mass >44 kg recorded from the Indo-Pacic region (tropical China to New Guinea) from
the Late Pleistocene to present day, showing the maximum geographical range, most recent time period when the species was
extant within the region, and estimated range of body masses. The table
underestimates
extinctions, since Late Pleistocene fossils
have usually been assigned to the nearest living relative or to a single extinct taxon, while recent molecular studies have tended
to divide extant taxa into geographically separated species. Almost all extant species occupy <10% of their maximum Holocene
range and an even smaller percentage of their Pleistocene one (Table 1 continues on page 120)

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Altered Ecologies: Fire, climate and human inuence on terrestrial landscapes120
Taxon Maximum range Last known extant Mass (kg) Common name
Helarctos malayanus China to Java extant 25-65 sun bear
Ursus thibetanus China to ailand extant 50-180 Asiatic black
bear
PERISSODACTYLA
Rhinocerotidae Dicerorhinus
sumatrensis
China to Borneo extant 500-1000 Sumatran
rhinoceros
Rhinoceros
philippinensis
Luzon Pleistocene <800
R. sinensis China, Vietnam after 40,000 BP >1000
R. sondaicus China to Java extant 1500-
2000
Javan rhinoceros
R. unicornis China to Java extant (S. Asia) 1600-
3000
Indian
rhinoceros
Tapiridae Megatapirus
augustus
China to Laos Holocene >500 giant tapir
Tapirus indicus China to Java extant 200-400 Malayan tapir
ARTIODACTLYA
Suidae Babyrousa spp. Sulawesi & islands extant 50-100 babirusa
Sus spp. China to Sulawesi extant 10-150 pigs
Cervidae Axis calamianensis Calamian Islands extant 30-90 Calamian deer
A. kuhlii Bawean extant 36-50 Bawean deer
A .porcinus China to ailand extant 36-50 hog deer
Elaphodus
davidianus
China recent 150-200 Père David’s
deer
Cervus eldii China to ailand extant <150 Eld’s deer
Rucervus
schomburgki
ailand 1932 100-120 Schomburgk’s
deer
Rusa alfredi Visayan Islands extant 25-80 Visayan spotted
deer
R. marianna Philippines extant 40-60 Philippine deer
R. timorensis Java &Bali extant 75-160 Javan rusa
R. unicolor China to Borneo extant 100-300 sambar
Bovidae Bos frontalis China to Malaysia extant 700-1500 gaur
B. javanicus China to Java extant 600-800 banteng
B. sauveli Vietnam to
ailand
recent <900 kouprey
Bubalus spp. Philippines late Pleistocene
B. arnee Myanmar-Java extant 800-1200 wild water
buffalo
B. depressicornis Sulawesi extant 150-300 lowland anoa
B. mephistopheles China mid Holocene <1000 short-horned
water buffalo
B. mindorensis Mindoro extant 180-300 tamaraw
B. quarlesi Sulawesi extant 150-300 mountain anoa
Pseudoryx
nghetinhensis
Laos, Vietnam extant 100 saola
Capricornis
milneedwardsii
China to ailand extant 50-140 Chinese serow
C. sumatraensis ailand to Java extant 50-140 Sumatran serow

Megafaunal extinctions and their consequences in the tropical Indo-Pacic 121
terra australis 32
New Guinea
e largest extant varanid in New Guinea (Varanus salvadorii) and the largest pythons (Morelia
spp.) may occasionally reach megafaunal size, and New Guinea supports two megafaunal
cassowary species (Wright 2005). e late Pleistocene fauna of the New Guinea highlands
also included several now-extinct marsupials of megafaunal size, including large browsing
kangaroos (Protemnodon spp.) and diprotodontids (Hulitherium, Maokopia, Zygomaturus)
(Long et al. 2002; Fairbairn et al. 2006).
Wallacea (excluding Sulawesi)
Potentially megafaunal reptiles on the oceanic islands of Wallacea (Figure 1) included varanids,
pythons and giant tortoises. Megafaunal varanids (Varanus komodoensis) are now confined to
Komodo, western Flores and a couple of nearby islands, and all giant tortoises are extinct, but
pythons (Python reticulatus) occur on most islands, apparently as natives. However, although
this species reaches megafaunal sizes on the mainland and continental islands, with a 6-7 m
snake weighing >50 kg (Fredriksson 2005), at least some of the Wallacean forms are dwarfed
(Auliya et al. 2002). Proboscids were apparently the only mammals able to swim the sea straits
east of Bali. Pleistocene fossil dwarf stegodons are known from several localities on Timor,
along with a giant tortoise and a large varanid, but all these seem to have vanished before the
first evidence for modern humans at 40,000 BP (O’Connor 2007). Fossil dwarf stegodons of
uncertain age have also been reported from Sumba (Sartono 1979) and from the tiny island
of Sangihe, between Sulawesi and Mindanao (van den Bergh et al. 1996). On Flores, a dwarf
(c. 300 kg) stegodon species from 900 ka was replaced by 850 ka by a larger species, which in
turn became dwarfed, before disappearing c. 12 ka (van den Bergh et al. 2008). e Komodo
dragon persisted throughout this sequence and is the only extant native megafaunal species,
but a giant tortoise was last recorded at 900 ka.
Figure 1. Map of Wallacea and the surrounding region

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Altered Ecologies: Fire, climate and human inuence on terrestrial landscapes122
Sulawesi
At the Last Glacial Maximum, the Makassar Straits separating Sulawesi from Borneo were only
45 km wide at the narrowest point. e Pleistocene fossil fauna had a number of extinct genera,
including giant tortoises, stegodonts, elephants and a large suid, Celebochoerus heekereni, but it
is not clear whether any of these survived into the late Pleistocene (van den Bergh et al. 2001).
e extant megafauna includes the babirusas, a pig (Sus celebensis), two species of anoa, and
the reticulated python.
Philippines
At Pleistocene low sea levels, most of the land area of the Philippines was in six major islands,
separated by narrow straits. Only Palawan and the adjacent small islands may ever have been
connected to Borneo and it is not clear whether or when this last happened. Palawan supports
a subset of the Bornean fauna, including species not found elsewhere in the Philippines, but
it also has several endemic species and subspecies. Tigers were present until at least the late
Pleistocene and a deer became extinct in the mid Holocene (Piper et al. 2008). e extant
Calamian deer is endemic to islands in the Palawan Group. Excluding the Palawan Group,
the extant megafauna of the Philippines consists of endemic pigs, deer and the tamaraw, plus
the widespread reticulated python (which is possibly a human introduction on some islands).
ere are also fossils of extinct Bubalus species on Luzon and Cebu, suggesting this genus may
once have been present throughout the Philippines (Croft et al. 2006). ere are poorly dated
Pleistocene fossils of stegodonts on Luzon and Mindanao, and fossils of both elephants and
rhinoceroses on Luzon (Bautista 1991; van den Bergh et al. 1996).
Java
e megafauna coexisting with H. erectus on middle Pleistocene Java included stegodons,
hippopotamuses and hyenas (Louys et al. 2007), but Java appears to have had a fully modern
fauna by 120 ka (Westaway et al. 2007), and – assuming that the fossils are correctly attributed
to modern species – no species recorded since then has become globally extinct. e late
Pleistocene megafauna consisted of the Asian elephant, orangutan, tiger, leopard, sun bear,
Malayan tapir, Javan rhinoceros, Javan rusa, banteng, wild water buffalo, Sumatran serow,
Javan warty pig, Eurasian wild pig and two species of python. e elephant, orangutan, tiger,
sun bear, tapir and serow disappeared in prehistoric to early colonial times and the Javan
rhinoceros has been reduced to a single population of 60 individuals. An extant endemic
species of deer occurs on Bawean Island, 150 km north of Java.
Borneo and Sumatra
e only fossil evidence of proboscideans from the huge continental island of Borneo is teeth
of uncertain provenance (Cranbrook and Piper 2007), while the current population of wild
elephants in the northeast seems best explained by the traditional story of introduction from
Java via Sulu (Cranbrook et al. 2008). A giant (probably megafaunal) species of pangolin
is known from parts of a single individual dated at c. 40,000 ka (Piper et al. 2007a). e
known Holocene megafauna included three taxa: Javan rhinoceros (Cranbrook and Piper
2007), Malayan tapir (Cranbrook and Piper 2009) and tiger (Piper et al. 2007b), which
became extinct before – possibly just before – modern times. e surviving megafauna
consists of the Bornean orangutan, sun bear, Sumatran rhinoceros, bearded pig, sambar deer,
banteng and reticulated python. Sumatra has a poorer fossil record than Borneo and the only
known megafaunal extinction to occur between the late Pleistocene and historical times is the
leopard (Louys et al. 2007), while the Javan rhinoceros is a recent extinction. In contrast to
Borneo, elephants, tigers and tapirs still survive, along with the Sumatran orangutan, sun bear,

Megafaunal extinctions and their consequences in the tropical Indo-Pacic 123
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Sumatran rhinoceros, bearded pig, sambar deer and reticulated python. All but pigs, deer,
tapirs and pythons are currently severely threatened by hunting (Corlett 2007).
Continental Southeast Asia and South China
During the middle Pleistocene, hominins in South China and Vietnam coexisted with
a diverse megafauna that included a giant (estimated 540 kg) ape, Gigantopithecus, which
became extinct around 300 ka (Rink et al. 2008). Some elements of this fauna persisted
into the late Pleistocene and several taxa that are now globally extinct survived into the early
Holocene. ese late survivors include at least one species of stegodon, a giant tapir and the
spotted hyena (Louys et al. 2007). Note, however, that these Holocene dates are considered
questionable by Turvey (2009a). In the middle to late Pleistocene, orangutans and giant
pandas occurred at many sites from southern China to ailand (Louys et al. 2007; Zhang et
al. 2007). e Malayan tapir is also recorded from late Pleistocene sites in China.
Megafaunal range declines and extinctions have been better documented in China than
elsewhere, with more archaeological sites and longer historical records. Stegodons, Malayan
and giant tapirs, hyenas and orangutans disappeared before the start of historical records. e
short-horned water buffalo (Bubalus mephistopheles) became globally extinct in early historical
times (Yang et al. 2008). e decline of the giant pandas started in the late Pleistocene and
accelerated in historical times (Loucks et al. 2001; Zhang et al. 2007), while the retreat of
the elephants and rhinoceroses (one to two species) has occurred largely over the past 3000
years, with a progressive withdrawal of the northern boundaries of their distributions in the
face of growing population pressures (Corlett 2007). Historical data suggest a threshold value
of about four people per square kilometres for rhinos, and 20 people for elephants, above
which they did not persist (Liu 1998). Rhinoceroses became extinct within the past 30 years
in China and elephants have been reduced to tiny populations near the southwestern borders.
e declines started later in continental Southeast Asia, but only a tiny population of Javan
rhinoceroses and a few scattered Sumatran rhinoceroses still survive, and the Asian elephant
occupies a fraction of its 19th century range (Corlett 2007). Tigers have been eliminated from
southern China and are in rapid decline everywhere else in the region, while leopards and the
two Southeast Asian bear species have been eliminated from most of the more accessible parts
of their continental ranges.
All tropical Asian bovids are currently threatened by hunting (Corlett 2007) and the
kouprey may be extinct. Deer are often considered relatively resilient to hunting, but all species
have declined severely in recent decades, including the widespread sambar, and this group
includes the only two global extinctions (in the wild) in the region within the past 150 years.
Schomburgk’s deer inhabited marshy grasslands in the central plains of ailand, but has
been extinct since 1938, while further north, Père David’s deer was eliminated from riverine
marshland in the lower reaches of the Yangtze River more than a century ago and recently has
been reintroduced. Eld’s deer, which occurs in open forests, has also had its range drastically
reduced over the past century. Pigs are the ‘last large mammal standing’ in many areas, but pig
populations have shown massive recent declines as a result of overhunting.
Why?
Many early and middle Pleistocene extinctions appear to represent a natural process of
species turnover, although there are some conspicuous exceptions, such as the giant tortoises
and hippopotamuses (Louys et al. 2007), but by the late Pleistocene, extinctions without
replacement predominate, matching the pattern seen in the Quaternary Megafauna Extinction
event elsewhere on the planet (Barnosky 2008). Unlike other well-documented examples of
the QME, however, there is no evidence to suggest a single, sharp extinction event, and no

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Altered Ecologies: Fire, climate and human inuence on terrestrial landscapes124
obvious peak with the presumed arrival of modern humans around 50,000 years ago. Instead,
there has been a steady trickle of regional then global extinctions, mostly within the past
40,000 years and many within the Holocene. e inadequacies of the fossil record and its
dating have inevitably blurred the true history of megafaunal extinctions in the region, but
surely not enough to account for the broad spread of apparent extinction dates.
e megafaunal losses described above have been attributed by different authors to various
combinations of direct (i.e. hunting) and/or indirect (habitat modification) human impacts
and natural environmental change. If the region had been completely uninhabited for the past
million years, it would be possible to build a plausible case for attributing most of the pre-
recent losses to environmental change, in particular the development of dense, closed-canopy
forest over all but the driest parts of the region at the start of the Holocene. Ground-dwelling
herbivores are denied access to the majority of the leaf production in a closed-canopy forest,
since this takes place in the canopy. A decline in the density of terrestrial herbivores in turn
threatens the largest carnivore, the tiger, which does not have access to arboreal prey, as well as
putting pressure on the leopard. e extinction of the giant pangolin may also reflect the loss
of open habitats that have a high density of accessible ant and termite nests at the start of the
Holocene (Medway 1972). A problem with such environmental explanations, however, is they
require that late Pleistocene environments and/or the transition to the Holocene were uniquely
difficult periods for megafaunal species. Each glacial and interglacial period is different, but
there is no independent evidence that the past 100,000 years has been significantly more
difficult for large vertebrates than previous glacial cycles (Barnosky et al. 2004). Volcanoes
have also been invoked as agents of extinction, particularly on Flores, where faunal turnovers
at 900 ka and 12 ka both coincided with volcanic eruptions. However, the largest eruption on
Earth during the late Pleistocene, that of Toba on Sumatra 74,000 years ago, did not coincide
with any known extinctions in the region (Louys 2007).
Environmental explanations that may be plausible in the absence of people become less
plausible in their presence. e capabilities of Homo erectus are largely unknown, but it is hard
to imagine a large, relatively intelligent, omnivorous, social primate coexisting for long with
something as vulnerable as a giant tortoise. Giant tortoises are known from Sulawesi, Flores
and other islands in the early to middle Pleistocene, and disappeared on Flores around the
time hominins, presumably H. erectus, arrived (van den Bergh et al. 2001). H. erectus was
associated at various times and places with other large vertebrates that no longer survive in
the region. A dwarf stegodon was lost from Flores at the same time as the giant tortoise, and
the bones of its dwarfed successor are associated with those of H. floresiensis in late Pleistocene
cave deposits (van den Bergh et al. 2008). Extinct megafauna, including elephants, stegodons
and rhinoceroses, are associated with early stone tools in the Philippines (Bautista 1991).
Human fossils and/or stone tools from middle and late Pleistocene cave sites in southern
China and Indochina are also often associated with the remains of stegodons and the giant
tapir (Ciochon and Olsen 1991; Bekken et al. 2004; Schepartz et al. 2005). Early Homo
was also associated at some sites with Gigantopithecus (Ciochon et al. 1996; Harrison et al.
2002). ese associations suggest some form of interaction, but there is no direct evidence for
hunting, and scavenging is equally plausible in most cases.
Although the evidence for an impact of H. erectus on megafaunal survival is weak, it is
surely significant that the clearest association between human arrival and megafaunal loss is in
the New Guinea highlands (Fairbairn et al. 2006; Field et al. 2008), which H. erectus did not
reach. e lack of a single, clear extinction event in tropical Asia could thus simply reflect the
complex chronology of hominin habitation. In contrast to most other regions of the world
outside Africa, where naive megafauna were faced by expanding populations of technologically
sophisticated, habitat-generalist, modern humans, tropical Asia’s fauna had a relatively gradual
introduction to the perils of humanity. If, as seems likely, H. erectus avoided closed-canopy
forest and, at least initially, lacked projectile weapons and mastery of fire, impacts may have

Megafaunal extinctions and their consequences in the tropical Indo-Pacic 125
terra australis 32
been slow and selective in comparison with those of modern humans, and may have allowed
time for some adaptive evolution.
With modern Homo sapiens, there is no need to speculate on its hunting ability, but it
is reasonable to question whether human population densities before the second half of the
Holocene were high enough to lead to faunal extinctions. e absence of a well-defined mass-
extinction event in tropical Asia in the 60,000-40,000 BP period during which modern humans
probably arrived is consistent with the ‘coastal express train’ model, with coastal populations
moving on as they depleted resources, and only later moving inland. e extinction of the
giant pangolin soon after the presumed arrival date was attributed above to the loss of open
habitats, but a slow-moving, 2 m long mammal whose only defence was to roll up must also
have been exceptionally vulnerable to hunting. A similar argument can be made for the slow-
moving, slow-breeding orangutans, which ranged widely over tropical East Asia in the late
Pleistocene, from southern China to Java, but were confined to the sparsely populated ever-
wet rainforests of Borneo and Sumatra by historical times (Delgado and van Schaik 2000).
e giant panda and the New Guinea megafaunal marsupials may also have been particularly
vulnerable to hunters for similar reasons, but there seems to be no obvious pattern to most
other pre-recent extinctions.
Many authors have combined the two major hypotheses and argued that a deteriorating
environment combined with pressure from hunting or anthropogenic habitat-modification to
push species over the edge into extinction. is is in many ways an unsatisfactory compromise,
and is hard to test or refute, but it is equally difficult to argue that either factor – drastic changes
in climate and vegetation, or the arrival and expansion of successive human species – had
no significant impact on megafaunal populations. A predominance of human impacts over
environmental change becomes increasingly clear over the past few thousand years and the
role of human exploitation in the drastic range reductions shown by all surviving megafaunal
species in the past century is undeniable (Corlett 2007).
Consequences
Whatever the causes of the large reductions in megafaunal diversity and biomass over the
past 130,000 years, the consequences are likely to have been, and continue to be, significant.
Indeed, if megafaunal impacts were significant, the vegetation and associated fauna of much
of the region may now be in a state of long-term relaxation from these impacts (Johnson
2009). Very large herbivores may have kept vegetation in a more open, patchy condition than
exists today. More generally, they may have acted as ‘ecosystem engineers’, modifying the
physical environment in a way that affects other species (Pringle 2008). e largest surviving
megaherbivores, the forest elephants and rhinoceroses, are browsers, with the strength to
push down, break off, or uproot shrubs, saplings and small trees (Corlett 2007). Elephant
movements can lead to the creation of extensive networks of trails. Loss of megafauna-
maintained open habitats would have impacted shade-intolerant plants and smaller terrestrial
herbivores that would have suffered reduced access to plant biomass. Large deer and cattle do
less incidental damage to vegetation, but are more selective as browsers and grazers. Plants that
have coevolved with these large herbivores may invest in physical or chemical defences that are
ineffective against smaller herbivores, and may thus lose out in competition with plants that
lack these defences when large browsers are removed (Johnson 2009). Dung piles from large
herbivores have multiple impacts, creating patchiness and providing habitat for other species
(Campos-Arceiz 2009).
In central Africa, forest elephants at natural densities probably disperse more seeds than
any other vertebrate species and disperse them over much larger distances (Blake et al. 2009),
but dispersal by megaherbivores has received little attention in tropical Asia. Many ripe, fleshy

terra australis 32
Altered Ecologies: Fire, climate and human inuence on terrestrial landscapes126
fruits reach the ground uneaten, where many are consumed by large terrestrial herbivores,
these herbivores including probably all extant species of pigs, deer, cattle, tapirs, elephants and
rhinoceroses (Corlett 1998). e indehiscent pods produced by some legumes are consumed
by the same animals. A ‘megafaunal syndrome’ of very large fleshy fruits (>10 cm diameter
with numerous small seeds, or 4-10 cm diameter with a few large seeds) dispersed largely
by mammals with >1000 kg body mass has been identified in Africa and Brazil (Guimaraes
et al. 2008), but the distribution of such fruits in the tropical Indo-Pacific has not yet been
documented. Asian elephants have been reported to prefer large, yellow, sweet-smelling fruits
with large, hard seeds (Kitamura et al. 2007). Some studies suggest that they consume less
fruit and fewer species than their African relatives, but fruits from 29 species were recorded by
mahouts in Myanmar as eaten by work elephants (Campos-Arceiz et al. 2008a), and potential
seed dispersal distances are very large (<6 km, Campos-Arceiz et al. 2008b). Rhinoceroses also
eat large fruits and can potentially move >10 km within plausible gut-passage times (Corlett
2009b). Tapirs have a similar potential.
e 44 kg cut-off used in this study brings in a wider range of fruit types than the classic
megafaunal syndrome described from studies on African elephants. Large deer disperse seeds
both by regurgitation of large, hard seeds from fleshy fruits (e.g. Prasad et al. 2006) and by
defecation of small ones, with some of the latter swallowed incidentally during consumption
of foliage (Myers et al. 2004; Yamashiro and Yamashiro 2006). Pigs and bovids disperse small
seeds in the same way as deer, but their role, if any, in dispersing larger seeds is not known.
e extinct megafaunal marsupials of New Guinea probably dispersed some seeds in the same
way as extant grazing and browsing megaherbivores (Webb 2008), but any role they had in the
dispersal of large seeds is probably covered by the extant cassowaries, which still disperse seeds
in large fruits (<6 cm) for long distances (Wright 2005; Bradford et al. 2008). Orangutans
are the largest arboreal frugivores and eat fruits of many different types, including the biggest
species available. eir ability to move large seeds over long distances (>1 km) is shared with
only a few other large-bodied frugivores (Corlett 2009b).
Megafaunal introductions
Many megafaunal species have been deliberately introduced by people outside their natural
ranges and a proportion of these have established wild populations. ese successful invasions
provide evidence for the existence of ‘empty niches’ in communities throughout the region,
particularly on islands, although in some cases these niches have been created or expanded
recently by human modification of the natural vegetation. Pigs (Sus scrofa, S. celebensis, Larson
et al. 2007) and deer (mostly Rusa timorensis) were spread throughout the region during the
Holocene. e archaeological record for Flores, for example, shows that the only megafaunal
animals at the start of the Holocene were people and Komodo dragons, but that people
subsequently introduced first the Sulawesi warty pig (Sus celebensis), c. 7000 years ago, and
subsequently the Eurasian pig (S. scrofa) and several smaller mammal species (van den Bergh et
al. 2008). e archeological record of south Sulawesi shows that deer (Cervus timorensis) first
appeared there about 4000 years ago (Simons and Bulbeck 2004). e elephant population
on Borneo is apparently also a recent introduction (Cranbrook et al. 2008).
Megafaunal reintroductions?
If the extinct megafauna played a unique role in the ecosystems they inhabited, it makes sense
to consider their reintroduction or replacement. Where megafaunal species have been lost
from an area in recent times, but persist elsewhere, then reintroduction is an option. Indeed,
many megafaunal species are likely to be favoured by the leaf and shoot biomass available near

Megafaunal extinctions and their consequences in the tropical Indo-Pacic 127
terra australis 32
ground level in disturbed and fragmented habitats. Piper and Cranbrook (2007) propose the
reintroduction of the Malayan tapir to the Planted Forest Zone in Sarawak, which consists of
490,000 ha of primary, secondary and industrial plantation forests, while several projects are
attempting to re-establish orangutans in areas from which they have been extirpated (Corlett
2009a). With an estimated 16,000 captive elephants in Asia and many of these unwanted as
their use in logging declines, the major problem with the Asian elephant is a lack of suitable
habitat, rather than a shortage of animals for release (Leimgruber et al. 2008). In contrast, both
species of forest rhinoceros are critically endangered and neither has a captive population.
Where the extinction was many tree generations ago, it is likely that the current vegetation
represents a new equilibrium in which previously suppressed plant species are more abundant
and others less abundant than when the megafauna was present at natural densities (Johnson
2009). While restoring the natural situation may seem a laudable aim, it is also possible that
the disruption of fragmented forest communities will simply promote invasive alien species,
since these are likely to exert a much greater ‘propagule pressure’ than any rare native species.
Where the missing taxon is globally extinct, but an ecologically similar relative persists,
then the introduction of this as a substitute could be considered. A major problem, however,
lies in assessing ‘ecological equivalence’. For proboscidians, the Asian elephant is the only
Asian survivor of a once diverse group. e fact that some of these other taxa coexisted with
Asian elephants in the past suggests that they are not complete equivalents, but how much this
matters is hard to determine.
Conclusions
e relatively small number of global extinctions in the Indo-Pacific megafauna in comparison
with the rest of the world masks a catastrophic collapse in local species diversity and biomass
since the middle Pleistocene. e evidence from the region is frustratingly incomplete, but
I consider it most likely that hominin impacts have been the major factor behind most large
vertebrate extinctions and range restrictions in the past 130,000 years and probably some
earlier ones. Large vertebrates have evolved in or invaded every accessible land mass on Earth
and appear to have been a major component of all vertebrate communities since soon after
vertebrates evolved, apart from the period immediately after the K/T extinctions 65 million
years ago. For their disappearance during the brief period following the expansion of the genus
Homo out of Africa to be a mere coincidence would require much stronger evidence than any
that is currently available for uniquely extreme environmental pressures during this time. Any
uncertainties about the role of human populations disappear by the mid Holocene, and the
devastating human impacts of the past century are well documented.
If an anthropogenic explanation for the megafaunal collapse is accepted as most likely,
this then raises the question of why there is no clear evidence of rapid overkill soon after
the arrival of humans. I have suggested a two-part explanation for this: that Homo erectus
lacked (or, at least, initially lacked) the technical skills and forest adaptations needed to cause
rapid population collapse, except in the most vulnerable of non-forest taxa, while H. sapiens
spread initially along the coast and invaded the forested interior only gradually and unevenly.
Indeed, much of the region had extremely low human population densities into historical
times (Corlett 2009a). Although the megafauna is equally absent whether it was wiped
out by climate change or by humans, the former would be an interesting palaeoecological
phenomenon, while the latter, if true, is a historical tragedy and one which we would reverse if
we could. Yet the surviving megafauna is everywhere in retreat and more extinctions are likely
(Corlett 2007). Conservation successes in the Indo-Pacific tend to be local and small scale,
while the larger-scale efforts needed to save viable populations of the largest species appear
to be beyond our current capabilities. Some species, such as tigers, will survive in captivity

terra australis 32
Altered Ecologies: Fire, climate and human inuence on terrestrial landscapes128
whatever their fate in the wild, but others, including the two critically endangered forest
rhinoceroses, lack viable captive populations. More than a billion people live in the region
covered by this review and this population will continue to rise until at least 2050 (Corlett
2009a). Conservation problems are many and urgent as a result, but the largest vertebrates
deserve special attention.
Acknowledgements
Geoff Hope was hugely influential in my early career, so he is the first person I need to
acknowledge. I have also benefited greatly from discussions with Ahimsa Campos-Arceiz.
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