Part II Species Accounts

Chapter
Sulawesi Babirusa Babyrousa celebensis

6 (Deninger, 1909)
Alastair A. Macdonald

Names
Genus: Babyrousa (Perry, 1811)
Species: Babyrousa celebensis (Deninger, 1909)
Names in other languages: French: Babiroussa des Célèbes; German: Sulawesi-Hirscheber; Italian: Babirussa di Celebes; Spanish:
Babirusa de Célebes; Minahasa, Sulawesi: Kalowatan.
The babirusa on the island of Sulawesi, Indonesia, have been the focus of local attention for about 35,400 years or more (Heekeren
1952, 1972; Eriawati 2003; Aubert et al. 2014). This is evidenced by the paintings of male and female images of these animals on the
walls of caves, East of Maros, in the southwest peninsula of the island (Figure 6.1A,B).
The fossil record is currently lacking. Briefly, four Pleistocene teeth found at Sompeh and Beru in South Sulawesi (Bartstra
1997) were initially described as Babyrousa babyrussa beruensis by Hooijer (1948). However, he subsequently reclassified these as
more probably Celebochoerus heekereni (Hooijer 1954). Although the current hypothesis is that the babirusa has developed since
Oligocene times (Thenius 1970), there is also the suggestion that the progenitor of the babirusa may have reached central Sulawesi at
a very much earlier date, perhaps during the Palaeogene (Van den Bergh et al. 2001). More recent, Holocene, sub-fossils of babirusa
have been found within the Tolian deposits (Sarasin & Sarasin 1905), collected from caves and rock shelters in south Sulawesi; these
have been described by Hooijer and named B. b. bolabatuensis (Hooijer 1950).
The babirusa is a somewhat bizarre pig in that the upper or maxillary canine teeth of the male grow through the skin of the nose
and curl over the face (Figure 6.2A). The growth process involved has recently been published (Macdonald et al. 2016). In the female,
these teeth are either markedly reduced or are absent. Attention external to eastern Indonesia was attracted to the animal in part by a
local story (Purchas 1625) reported in about 1512 to the Portuguese sea captain Antonie Galvano by the ‘King of Tydore’; he was told
that on the ‘Islands of Batochina . . . there are hogs also with horns’.
59

Published online by Cambridge University Press

 Part II: Species Accounts

Figure 6.1  (A) Rock-painting, in red striped-line technique, of a charging (babirusa) boar (Heekeren 1972, plate 59). (B) Drawing of the charging (babirusa) boar in
(A) (Heekeren 1972, p. 117).

Figure 6.2  (A) Adult male babirusa from north Sulawesi. Note the dark-coloured wet areas of his body revealing that he had recently been wallowing in mud.
© M. Patry. (B) Adult female babirusa with suckling young from north Sulawesi. Note that the mother is ingesting warm mud from between the stones. © M. Patry.

Taxonomy Descriptive Notes

A group of 10 skulls from the eastern part of the northern penin- Babyrousa celebensis is characterized by its sparse body hair and
sula of Sulawesi and the adjacent island of Lembeh was described a nearly hairless tail tuft. The adult male head and body length
by Deninger (1909) as different from the skulls from the island is 85–110 cm, with a shoulder height of 60–65 cm. Their body
of Buru. He assigned the name ‘Babirusa Celebensis’ to them. weight may reach 100 kg (Figure 6.2A). The long, relatively thick
The local people called these animals Kalowatan (Macdonald maxillary canines of the adult male emerge through the skin of
and Leus, unpublished). By association, and for want of data in the nose and curl dorsally towards the forehead. The canine
2008, all living babirusa on mainland Sulawesi were similarly teeth of female babirusa are very small or absent (Figure 6.2B;
called B. celebensis (Macdonald et al. 2008a). The taxonomy of Deninger 1909; Mohr 1960).
the babirusa is currently being re-examined (Macdonald et al.,
unpublished). Habitat
Today, babirusa inhabit tropical rainforest, and are active during
Subspecies and Distribution daylight hours (Patry et al. 1995; Clayton & MacDonald 1999).
The babirusa that once lived in the south-west peninsula are Whereas previously the animal had been reported to occur in
represented by sub-fossil remains described by Hooijer (1948, low-lying areas near coasts, recent anecdotal and survey reports
1950, 1954). The current distribution is shown in Figure 6.3. indicate that the species is now confined mostly to the interior,
60

Published online by Cambridge University Press

 Chapter 6: Sulawesi babirusa Babyrousa celebensis (Deninger, 1909)

Figure 6.3  Sulawesi babirusa distribution (Source: IUCN 2016, Red List of Threatened Species). (A simplified black and white version of this figure will appear in some
formats. For the colour version, please refer to the plate section.)

on higher and less-accessible ground. There the tree cover is pre- babirusa will normally travel, but fragments of information sug-
dominantly tropical lowland evergreen and semi-evergreen rain- gest that it is quite restricted. Because it was impossible to dif-
forest. These are not dominated by any particular family of trees ferentiate between the females entering the salt-lick study sites,
(Whitmore et al. 1989; Whitten et al. 2002). Palms are common it was not possible to say whether these females had remained in
in the lowland forest, including Oncosperma horridum, Liculala the same locality or had a wide range that overlapped with those
celebensis, Pinanga, Areca, Caryota, and Livistona rotundifolia of other females (Patry et al. 1995; Clayton & MacDonald 1999).
(Whitten et al. 2002). Ebonies (Diospyros spp.) were also com- However, because the males could be identified from one another
mon in dense clumps. The dipterocarps included Anisoptera cos- by the shape and size of their canine teeth (Figure 6.2A), the
tata, Hopea celebica, H. gregaria, Shorea assamica, Vatica rassak, observations suggest that the adult male babirusa do have home
and V. flavovirens. Many different kinds of fruit-bearing trees ranges, and that these overlap at the salt-lick with those of a num-
are distributed throughout these forests. Although the detailed ber of other adult males. It is possible, therefore, that home ranges
nature of the babirusa’s association with these various species of may not be large where they permit sufficient access to food and
food-bearing plants has not yet been studied, those reports that water, salt-lick components, and members of the opposite sex.
have been published were gathered together and analysed by Leus There is evidence that babirusa on Sulawesi can swim rela-
(1994, 1996). They included Ficus spp., Canarium sp., Elatostema tively large distances: a single adult male babirusa was reported
sp., Lithocarpus sp., Mangifera sp., and Pangium edule. approximately 500 m from the shore of Lake Poso in Sulawesi,
across which it appeared to be swimming (Melisch 1994). This
observation substantiates claims frequently repeated in the
Movements and Home Range older literature. Recent experience of babirusa in several zoo-
logical gardens reinforces the understanding that babirusa
Territorial Range derive benefit from lying in and swimming through water.
Very little is known about the territorial ranges used by babirusa The skeletal and muscular anatomy of the babirusa that ena-
in the wild. Nothing has yet been reported to indicate how far bles terrestrial and aquatic locomotion has been reviewed and
61

Published online by Cambridge University Press

 Part II: Species Accounts

described (Kneepkens et  al. 1989; Macdonald & Kneepkens morning. This behaviour did not involve rooting if the ground
1995; Kneepkens & Macdonald 2010). was dry or compact because the nose of the babirusa lacks a
It is clear that babirusa mark territory. Studies in zoologi- large rostral bone. In other pigs this structure provides sup-
cal collections have shown that adult male babirusa do this in port for the tough connective tissue plate of the rhinarium. In
a variety of ways (Leus et al. 1992, 1995). The side of the face is its place the babirusa has a thin, partially calcified membrane
rubbed on tree saplings, branches and other uprights, such as instead (Macdonald, unpublished). Food items were looked
posts or fence rails in the pen. The male will also plough with its for on the ground, under logs and stones, in wet areas and on
nose in soft sandy soil, apparently depositing secretions from bushes and trees (Leus & Vercammen 1996). Animals would
the face and salivary glands in the substrate. In the forests of stand with their forelimbs against tree trunks to reach hanging
North Sulawesi, adult males have been observed face-rubbing leaves and fruit. They could even stand free of such support on
on the stems of forest saplings, branches and small trees; plough- their hind limbs to pluck leaves from the trees (Leus et al. 1995;
ing behaviour in soft substrate has also been described (Patry Macdonald & Leus 1995). From about mid-morning onwards
et al. 1995). The babirusa male has also been filmed marking the an increasing proportion of the time was taken up with non-
ground adjacent to wallows, by rubbing the side of his face on foraging activity; the animals were either wallowing or lying
the substrate (Macdonald et al. 1996). Scratches on the canine down. In the wild, the numbers of animals visiting the salt-lick
teeth of museum skulls had earlier been mistakenly thought were seen to increase from mid-morning onwards (Clayton &
to be due to fighting behaviour (MacKinnon 1981). It has now MacDonald 1999). Both in the wild and in zoological collec-
been shown that it is ‘scent marking’ face-rubbing behaviour tions, males have generally been observed to wallow more often
that causes this very specific pattern of wear of the canine teeth. than females (Figure 6.2A; Leus et al. 1995).
Further studies are required to define the chemical nature of
the ‘scent marking’ molecules used by the babirusa (Macdonald
et al. 1996). Feeding Ecology
Food Supply and Digestion
Activity Patterns Babirusa are omnivorous; in zoos, adult babirusa have been
observed to chase, catch and eat small mammals and birds,
Daily Activity implying that they may show similar behaviour in the wild (Leus
Babirusa build nests in the wild (Guillemard 1886). These are et al. 1995). From observations of both wild and captive indi-
constructed from branches or leaves, and provide shelter from viduals they are known to consume a wide variety of leaf, root,
the rain and cold. Sleeping nests tend to have little or no padding and fruit material (Leus 1994, 1996; Macdonald & Leus 1995;
on the ground, and are essentially ‘babirusa-sized depressions’. Reksowardojo 1995). Studies suggest that fruits in particular
Nests built in the forest by sows for farrowing purposes were may be an important component of the diet (Leus 1994, 1996;
up to 3 m long and 25 cm deep, and were layered with branches Leus et al. 2001). Leus compiled an annotated list of species of
torn from trees and bushes (Macdonald 2008). Babirusa make trees and palms that grow in Sulawesi and produce fruits that
similar constructions of bedding in zoological collections, pro- may contribute to the diet of babirusa (Leus 1996). The dietary
vided they are supplied with suitable plant material (Melisch benefits gained by babirusa that ingest wet mud from warm salt-
1994; Leus et  al. 1995). When groups of babirusa have been licks have not been investigated in any detail (Patry et al. 1995;
penned together and have access to nest-building materials, all Clayton & MacDonald 1999; Leus et al. 2001).
the animals within the pen contributed towards the construc- Babirusa jaws and teeth are strong enough to crack very
tion of a communal nest, which they then shared overnight hard nuts with ease (Peters 1985; Leus et al. 1995). Seeds of a
(Macdonald 2008). Further studies are required, in the wild, to wide variety of fruits, including the canari (Canarium), oaks
discover whether communal nests are constructed by groups (Lithocarpus), figs (Ficus), and chestnuts (Castanopsis), are
of babirusa. Where no bedding has been provided in zoos, it is available on Sulawesi (Whitmore et al. 1989; Leus 1996). Fruits
noteworthy that babirusa slept together on the ground in groups are also an important component of the diet of Sulawesi Macaca
(Macdonald 1991; Leus et al. 1995). spp. The extent to which babirusa feed on the fruit pieces dis-
The daily activity profile of the babirusa has not been stud- carded by these primates also awaits study. The babirusa’s appar-
ied in the wild. However, some preliminary indication may be ent requirement for fruit-bearing trees, as a component part of
suggested by the distribution of daily behaviour exhibited by their environment, has been implied (Whitten et al. 2002); as
36 babirusa that were housed together as one large group in soon as mature trees were cleared by forestry operations or lands
Surabaya zoo (Leus et al. 1995). Animals slept at night, which was converted to food crop production, the babirusa were no longer
consistent with the general lack of activity at night recorded in to be found in that area. Further detailed studies, of the diet in
North Sulawesi (Patry et al. 1995; Clayton & MacDonald 1999). the wild and the fruiting patterns of the forest, are required to fill
About an hour before sunrise, individuals began to wake up these gaps in our current knowledge. They would help to explain
and move about. Voiding of faecal material and urine occurred how diet availability in wet and dry seasons might influence
at this time, and animals appeared to prefer to defecate under babirusa group size, structure and home range.
branches and tree trunks. Foraging extended throughout The anatomical and histological structures of the stom-
the day, but occupied a larger proportion of the time in the ach and digestive organs of the babirusa have been described
62

Published online by Cambridge University Press

 Chapter 6: Sulawesi babirusa Babyrousa celebensis (Deninger, 1909)

Figure 6.4  (A) Diagramatic representa-

tion of a ventral view of the stomach
of an adult babirusa with the relative
surface areas of the gastric glands and
the cardiac gland mucosa indicated. o,
oesophagus; p, pyloris. Modified after
Leus et al. (2004). (B) Scanning electron
micrograph of the unique honeycomb
appearance on the surface of the cardiac
gland mucosa, with a section to illustrate
the thin-walled tubes above each glan-
dular pit. Modified after Macdonald et al.
(2008b). Scale bar = 0.2 mm.

(Leus 1994; Leus et al. 1999, 2004; Agungpriyono et al. 2000; discharge (Leus et al. 1995). Oestrus usually lasts 2–3 days, and
Macdonald et  al. 2008b; Adnyane et  al. 2010). The stomach the female is not receptive to males at other times (Macdonald
has an elongated shape and a bulky appearance (Figure 6.4A). 1993). In zoological collections, females generally re-cycle
A blind sac with a diverticulum at its extremity is located to within 3 months after the birth of their young.
the left of the oesophageal entrance. The total luminal surface Studies have been undertaken on the anatomy and histol-
area of the adult stomach is about 3000 cm2 of which the car- ogy of the female and male reproductive tracts of the babirusa
diac gland area represents the largest portion (>70 per cent). (Berger et  al. 2006; Agungpriyono et  al. 2007; Ziehmer et  al.
This region was typically characterised by the ‘honeycomb’ pat- 2010, 2013). Their appearance is as illustrated in Figure 6.5.
tern of its luminal surface (Figure 6.4B). The thin-walled tubes, In the former, the thin-walled infundibulum almost com-
between 200 and 260 μm in length, that form the ‘honeycomb’ pletely encloses the ovary (Figure 6.5A). The uterus is bicornuate
are each sited over the pit of a cardiac gastric gland and are filled and relatively short in contrast to the relatively long cervix. The
by a stream of its secretion. latter is thick-walled and has a narrow, spirally arranged lumen.
Within that secretory mileu was found a complex bacterial Each testis lies in the scrotum that is located in a sub-anal position
microflora of rods and chains of cocci. The identities of most against the caudal surface of the thigh musculature (Figure 6.2A).
of these microorganisms remain unknown. However, the close The prostate, comprising a corpus and disseminate parts, lies
association they exhibited with the mucus secreted by the cardiac ­ventral to the vesicular glands partly embedded in the dorsal wall
glands indicated that this large area of babirusa stomach might of the urethra (Figures 6.5B,C). The ‘corkscrew-like’ twist at the
have been structured to form a stable and yet self-­refreshing envi- distal end of the penis corresponds to the shape of the cervix in
ronment for the housing and multiplication of what appear to be the female.
autochthonous commensal bacteria. The micro-­environment of The body weight of the sow increases during gestation
the cardiac gland area may be consistent with specialised micro- (Chaudhuri et al. 1990). The gestation length is usually 155–158
bial fermentation of ingesta. Alternatively, it may support micro- days, although up to 171 days has been reported (Vercammen
bial modification of glandular secretions. Several experiments 1991). The normal litter size is one or two uniformly brown-
have investigated food digestion by the babirusa (Conklin et al. coloured piglets, but a low incidence of triplet births has been
1994; Leus 1994, 2000; Leus & Macdonald 1997; Van Wees et al. recorded both in zoological collections and in the wild. One
2000; Clauss et al. 2008). Much research remains to be done on instance of four eggs having been shed from the ovaries of a zoo
this topic, however. animal has been reported (Ziehmer et al. 2013) and this matches
the report of a litter of four fetuses in utero in a wild female (Patry
1990). The babirusa has a diffuse epithelio-chorial placenta
Reproduction and Growth (Macdonald 1994).
Babirusa may become sexually mature as early as 10 months of Neonatal babirusa are small in size and weigh less than 800 g
age, with the age of sexual maturity in the wild being influenced at birth. There are usually four functioning mammary glands in
by a number of factors including the level of nutrition (Ogle & the babirusa, two located inguinally and the second pair on the
Macdonald 2008; Ziehmer et  al. 2010, 2013). Animals are abdomen; however, an additional pair of non-functional teats,
unlikely to breed until they are more than one year old. Indeed, slightly cranial to the others, has sometimes been reported (Leus
the earliest recorded pregnancies of babirusa in zoological col- et  al. 1995). Various aspects of nursing behaviour have been
lections occurred in animals aged about 18 months (Chaudhuri described (Bowles 1986; Patry et al. 1995; Maclaughlin 2000).
et al. 1990). Oestrus cycle lengths of between 35 and 37 days The composition of the milk (Chaudhuri et al. 1990) is reported
have been described by endocrine studies (Chaudhuri et  al. as: total solids = 23%; fat = 13%; protein = 7.8%; sugars = 2.7%.
1990; Berger et al. 2006). During the oestrus period, the external Suckling (Figure 6.2B) lasts for about 1–3 minutes.
labia of the vulva swell to twice their normal size and slightly In zoological collections, sows produce young at all times
evert, exposing the internal mucous membrane; this is associated of the year, and may produce two litters within a 12-month
with reddening of the vulvar area and the presence of a fluid period (Chaudhuri et al. 1990; Ogle 2010). However, as it seems
63

Published online by Cambridge University Press

 Part II: Species Accounts

Figure 6.5  (A) Diagramatic representation of the dorsal view of the non-pregnant female reproductive tract of the babirusa, with sectional views of the cervix,
vagina, and vestibule. Modified after Ziehmer et al. (2010). o, ovary; i, infundibulum; t, uterine tube; h, uterine horn; b, body of uterus; c, cervix; u, external urethral
orifice; v, vagina; w, wall of vestibule; g, glans of clitoris; l, lips of vulva. Scale bar = 20 mm. (B) Diagramatic representation of the dorsal view of the male reproductive
tract of the babirusa. Modified after Ziehmer et al. (2013). (C) Diagramatic representation of the left lateral view of the male reproductive tract of the babirusa. The
prostate gland lies between and ventral to the paired vesicular glands, and is partly embedded in the wall of the urethra. Modified after Ziehmer et al. (2013). (B,C) a,
bladder; b, vesicular gland; c, position of the prostate gland; d, bulbourethral gland with Musculus bulboglandularis; e, pelvic urethra; f, Musculus bulbospongiosus;
g, bulb of penis; h, (extended) sigmoid flexure of the penis; i; body of the penis; j, penis spiral; k, glans penis; l, left testis; m, head of epididimis; n, tail of epididymis; o,
ductus deferens. Scale bars = 20 mm.

likely that diet or seasonal factors would normally influence

inter-birth intervals, litters in the wild may be produced less Behaviour
frequently. Observation of babirusa in North Sulawesi revealed
adult females accompanied by sub-adult animals as well as juve- Social Behaviour
niles (Table 6.2; Patry et al. 1995). It was not clear whether these Babirusa are social animals (Macdonald et al. 1989; Patry et al.
youngsters of two different ages represented the progeny of con- 1995; Clayton & MacDonald 1999). Groups or troops of up
secutive litters from the same adult female. However, it has been to 13 animals have been observed in the rainforest, especially
observed in a zoo environment that there can be a close bond around water, communal wallowing areas, and warm salt-licks.
between adult female babirusa and their young that may last Aggregations as large as 46 individuals, comprising several dis-
for some time after weaning. It was also noticeable that females tinct groups, have sometimes been observed congregated at a
that were normally quite docile in zoological collections became salt-lick. One of the studies of babirusa in northern Sulawesi
exceedingly aggressive to their keepers and other babirusa, from (Patry et  al. 1995) enabled the following observations to be
shortly before parturition until about two weeks after the young quantified (Tables 6.1 and 6.2).
were born (Reinhard & Frädrich 1983; Leus et al. 1995). Studies Most sightings of solitary animals (84 per cent of all s­ olitary
still remain to be carried out in the wild to establish the precise animals) were of adult males. Single adult males were seen
nature of the sow’s behaviour at the time of parturition, and the with single adult females, but never with two or more females
relationship she has with other adult babirusa during the deliv- unless young animals were present (Table 6.2). Pairs or trios
ery and nursing of the young. Experience of small, mixed-sex of males were rarely seen, and bachelor groups of four or more
groups in some zoological collections suggest that it was not adult male babirusa have not yet been reported. Adult females
essential for the survival of the young for the pregnant female to were rarely seen without company; they were sometimes found
be on her own. In other zoological collections with similar-sized together with other adult animals, but most often they were with
groups, cannibalism of newly born animals has been experi- young babirusa (Figure 6.2B). Groups containing adult females
enced despite considerable attempts at the defence of the neo- and young numbered 84 (of the 226 groups studied), of which
nate by the post-parturient sow. ­two-thirds (56 groups) had no adult males present.
64

Published online by Cambridge University Press

 Chapter 6: Sulawesi babirusa Babyrousa celebensis (Deninger, 1909)

Table 6.1  The number of babirusa in each group (group size) and the These can be categorised as ‘threat at a distance’, ‘surprise rush’,
number of groups of this size seen visiting salt-licks in North Sulawesi.
‘nose in the air’, ‘head under jaws submission’, ‘front half sup-
Group size Number of groups ported’, and ‘boxing’ as the level of threat increased. The boxing
behaviour was the final stage observed, and this involved the two
1 104
adult males raising their forelimbs off the ground until both were
2 27 standing on their hind limbs facing one another (Figure 6.6A,B).
3 35 Once in this position the babirusa leaned and paddled
4 20 against the chest and shoulders of his opponent (Figure 6.6A).
5 15 Their snouts were held as high as possible (Figure 6.6B). If
one of the animals fell onto all four legs it reared up again and
6 12
the boxing continued. The boxing match usually lasted for
7 7
2–5 minutes, but could extend for up to 20 minutes. If the infe-
8 6 rior animal submitted at the earlier stage of ‘head under jaws
submission’ it would utter a very short squawk or a continu-
Table 6.2  Numbers of groups of babirusa, according to the number of
ous rattling stream of sound, the pitch and intensity of which
adult males and females and the presence or absence of youngsters, seen increased, as it seemed to perceive increased threat. Ironically,
visiting salt-licks in North Sulawesi. the rounded upper canine teeth of the positionally and socially
Adult males inferior male appear to function as protection to the throat of
the superior male, preventing potential (accidental) damage
0 1 2 3
from the lower canines of the inferior animal (Macdonald et al.
Adult females 1993; Leus et al. 1995). The teeth were never involved in ‘inter-
0 female only – 87 5 2 locked tooth’ wrestling matches (Blouch 1990; Macdonald et al.
0 female + young – 6 0 0 1993; Patry et al. 1995). The relatively shallow tooth sockets sup-
1 female only 6 7 2 1 porting the upper canines are not strong enough to withstand
such activity (Macdonald et al. 1993).
1 female + young 38 11 2 0
Female babirusa exhibit the ‘threat at a distance’ and ‘sur-
2 females only 3 0 0 0
prise rush’ agonistic behaviours, although ‘nose in the air’ and
2 females + young 13 7 4 0 ‘head under jaws submission’ were rarely seen between adult
3 females only 1 0 0 0 females, and ‘front half supported’ and ‘boxing’ were never seen.
3 females + young 5 2 1 0 When two females confronted one another their behaviour was
4 females only 0 0 0 0
often quite violent, fast-moving and noisy (Macdonald et  al.
1993; Patry et al. 1995). The objective seemed to be to bite the leg
4 females + young 0 1 0 0
or foot of the opponent.

Agonistic Behaviour Parasites and Diseases

Studies of agonistic behaviour in zoological collections and in Veterinary observations and autopsy reports from zoological
the wild have shown that adult males progress through a reper- collections indicate that the morbidity and mortality of older
toire of such behaviours (Macdonald et al. 1993; Leus et al. 1995). babirusa include several different tissue cancers. Little is known

Figure 6.6  Two frames captured from

the early morning jousting bout between a
pair of young adult male babirusa brothers
in Surabaya zoo. (A) The ‘boxing’ agonistic
behaviour is demonstrated. (B) The winner
of this ‘boxing’ contest extended his nose
above the head of his opponent who was
already lowering his head below the chin of
the superior animal. Photographs by Alastair
A. Macdonald.

65

Published online by Cambridge University Press

 Part II: Species Accounts

about the susceptibility of the babirusa to natural or introduced live longer in a zoo environment, and one female babirusa is
pig-borne diseases (Munro et  al. 1990). Hamerton reported a recorded to have lived for 24 years (Mohr 1960).
case of mass infestation of the lungs of a young babirusa with
Taeniidae (Echinococcus) cysticerci shortly after it arrived in Conservation Measures
London zoo from Sulawesi (Hamerton 1931). The diseases to
Opportunities are available to provide conservation sup-
which babirusa in zoological collections have been shown to be
port for the babirusa. The species has been accorded national
susceptible include the foot and mouth disease virus (Aphthae
protection under Indonesian law since 1931. It has been
epizooticae) in four male babirusa (Urbain et al. 1938), the bacte-
included on Appendix I of CITES since 1982. It has also been
rial Mycobacterium bovis disease which caused tuberculosis in a
categorised as ‘Vulnerable (VU – A2cd C1)’ in the IUCN Red
female (Hamerton 1941), and a fungal disease resulting in cuta-
Data Book (Macdonald et  al. 2008a; Leus et al. 2016), and
neous aspergillosis of a male (Lopez Gonzales & Taylor 1998).
listed as ‘Endangered’ by the United States Department of the
Interior (United States Department of the Interior 1980). It
Status in the Wild is currently protected by Indonesian law; Undang-un dang
Republik Indonesia Nomor 5 Tahun 1990 Tentang Konservasi
Threats to Survival Sumber Daya Alam Hayati Dan Ekosistmnya (Act No. 5/1990
Adult babirusa appear to have few, if any, natural predators. The Conservation of living resources and their ecosystems). In 2013,
small numbers of young born to the babirusa suggest that this the Indonesian government published a taxon-specific con-
animal is not adapted to a high rate of natural predation. No stud- servation strategy and action plan (Strategi Dan Rencana Aksi
ies seem to have quantified the predatory impact on babirusa of Konservasi Babirusa Tahun 2013–2022) (DKKH 2015). The
the larger snakes of Sulawesi. However, hunting by humans with national action plan identifies 11 priority conservation sites for
snares, nets, spears, and dogs has undoubtedly been an impor- babirusa on Sulawesi (Bogani Nani Wartabone Connected Area,
tant factor since prehistoric times (Hooijer 1948; Heekeren 1952). Sojol–Nantu Mountain Connnected Area, West and East Coast of
Studies over the last 40 years consistently show that widespread the narrow portion connecting North and Central Sulawesi, Lore
snare trapping constituted the main threat to the remaining popu- Lindu Connected Area, Morowali, Bakiriang Connected Area,
lations of babirusa in North Sulawesi (Blouch 1990; Clayton et al. Lombuyan, Latimojong Mountains, Takolekaju Mountains,
1997; Lee et al. 2005), and more recently, elsewhere in Sulawesi Verbek Mountains, and the Tangkeleboke–Abuki–Mataromba
(Milner-Gulland & Clayton 2002; Macdonald, unpublished). In Mountain Connected Area) (Indonesian Ministry of Forestry/
the past, the babirusa represented a by-catch to the hunting of the IUCN 2013). Memoranda were signed by the zoological associa-
Sulawesi warty pig (Sus celebensis). This would no longer appear tions in Indonesia, Europe, and North America to work together
to be true. The same prices are paid for the meat of both species, for the conservation breeding of the babirusa in the wild and in
which gives the hunters no incentive to selectively catch one pig zoological collections (Indonesian Ministry of Forestry/IUCN
in preference to the other (Clayton et al. 1997; Milner-Gulland & 2013). Relatively small populations of babirusa reportedly still
Clayton 2002). In south-east Sulawesi the babirusa is a by-catch of occur in a number of the national parks, nature reserves, hunting
anoa hunting by Muslim communities and is regarded as a ‘con- reserves, and protected forests in Sulawesi (Burton & Macdonald
taminant’ of the traps (Macdonald, unpublished). These babirusa 2009; Indonesian Ministry of Forestry/IUCN 2013). Discussions
are killed and their carcasses discarded in the forest. In other areas are currently underway to explore how best to promote their
the babirusa is a specific catch; they are to provide canine teeth to conservation within these areas. Education efforts supported
traders from the island of Bali who supply their use to the making by schools, newspapers, zoological societies, local and interna-
of Balinese topengs (dance masks) (Macdonald, unpublished). tional NGOs have distributed wildlife conservation information
In these various ways the population of babirusa in the wild to the children and raised awareness among the general pub-
throughout Sulawesi is being placed under severe and increas- lic on Sulawesi (Maklis 1991; Milner-Gulland & Clayton 2002;
ing market pressures. The widespread lack of public awareness Indonesian Ministry of Forestry/IUCN 2013). More is needed,
and understanding of the problem has allowed most of north- however. Recent anecdotal reports indicate that babirusa remain
east Sulawesi to be stripped of babirusa, with a permanent loss subject to local hunting. Pressure from an expanding human
to the genetic variability of the whole population that these ani- population and insufficient up-to-date and widely distributed
mals represented. information seem to combine with other factors to make much of
Large commercial and local-scale logging operations also the protective legislation ineffective at a local level. The amount
pose a major and increasingly serious threat to this species of suitable habitat remaining within the forest-degraded ‘pro-
(Smiet 1982; Abood et al. 2015). The loss and degradation of tected’ areas is less than that suggested by the map-designated
habitat has already resulted in the dramatic diminution in the sizes of these reserves. Small populations of babirusa isolated in
known range of this species in North and South Sulawesi, and relatively small pockets of forest are not viable in the medium
on the island of Sulabesi. Babirusa are one of the first animals to to long term (Frankham et  al. 2010; Indonesian Ministry of
become locally extinct after logging or land opening, and does Forestry/IUCN 2013). Efforts have been made to identify the
not appear to be as adaptable as the Sulawesi warty pig. range of genetic variability that still exists within the babirusa
It is not known how long babirusa live in the wild, but extrap- population (Macdonald et al., unpublished); forthcoming pub-
olating from the experience of animals in zoological collections, lications will inform both local and international conservation
it may be about 10 years (Ziehmer et al. 2013); animals tend to breeding before the animals it represents are killed off.
66

Published online by Cambridge University Press

 Chapter 6: Sulawesi babirusa Babyrousa celebensis (Deninger, 1909)

Status in Captivity
There are currently (15 December 2015) 77,113 = 190 babirusa
in 34 zoological collections worldwide (Figure 6.7). These are
managed regionally by studbook-keepers in Indonesia, Europe,
and North America. A Global Species Management Plan
(GSMP) (in conjunction with GSMPs for Banteng and Anoa)
for babirusa has been prepared during a workshop, from 25 to
30 January 2016, at Taman Safari Indonesia, in collaboration
with the Indonesian Ministry of Environment and Forestry and
the Indonesian Zoo Association. The application for the GSMP
was approved by the World Association of Zoos and Aquaria
(WAZA) on 16 March 2016 and a Masterplan (actions docu-
ment) is currently being written up. A GSMP committee has been
established, representing all relevant regional zoo associations
(including the Indonesian Zoo Association), the Indonesian
Ministry of Environment and Forestry and the IUCN Wild Pig
Specialist Group. The committee also includes an international
studbook keeper, a population biologist and in-situ experts. An
executive summary and set of recommended actions were pro-
duced from the workshop. A set of priority in-situ and ex-situ
actions has been agreed for the coming two years. Primarily,
these actions cover a global breeding and transfer plan for babi-
rusa, capacity-building for Indonesian zoos and government
authorities rescuing animals, support for education and aware-
ness programmes in zoos and the wild, and a range of in-situ
projects (currently being designed). Support mainly stems from
individual zoo institutions, and may be financial, in kind, or in
the form of direct advice and training given by zoo staff.

Acknowledgements
I should like to acknowledge the hospitality of Ulla Freifrau von
Mengden and the Utomo family in Jakarta, ‘Pak Yah Yah and Figure 6.7  Babyrousa celebensis at Taman Safari, Bogor (Photo by R. Wirth).
Mama Dios on Sulawesi, and the directors and staff of Surabaya (A black and white version of this figure will appear in some formats. For the colour
version, please refer to the plate section.)
zoo, Ragunan zoo and Taman Safari Indonesia I and II during
my visits to Indonesia. Information gathered from Sulawesi
would not have been possible without the assistance of my reported in this paper were derived from research on babirusa
colleagues, Dr Kristin Leus, the late Monsieur Maurice Patry, from Sulawesi, supported in part by the University of Edinburgh,
Professor Srihadi Agungpriono, and Bapak Hali Day. I am also the University of Edinburgh Development Trust, the Carnegie
glad to acknowledge the anatomical material, carefully curated Trust for the Universities of Scotland, the Royal Zoological
in museums in SE Asia, North America and throughout Europe, Society of Scotland, International Scientific Support Trust Ltd
to which I have been given access. The personal observations and the Balloch Trust, to all of which I am also most grateful.

References
Abood, S. A., Lee, J. S. H., Burivalova, Z., Agungpriyono, S., Macdonald, A. Aubert, M., Brumm, A., Ramli, M.,
Garcia-Ulloa, J. & Koh, L. P. A., Leus, K. Y. G., et al. (2000). et al. (2014). Pleistocene cave art
(2015). Relative contributions Immunohistochemical study on the from Sulawesi, Indonesia. Nature
of the logging, fiber, oil palm, and distribution of endocrine cells in the 514: 223–227.
mining industries to forest loss in gastrointestinal tract of the ­babirusa, Bartstra, G.-J. (1997). A fifty years
Indonesia. Conservation Letters Babyrousa babyrussa (Suidae). commemoration: fossil vertebrates
8: 58–67. Anatomia Histologia Embryologia and stone tools in the Walane valley.
Adnyane, I. K. M., McDonald, A. A. 29: 173–178. Quartär. Internationales Jahrbuch zur
Winarto, A. & Agungpriyono, S. Agungpriyono, S., Kurohmaru, M., Erforschung des Eiszeitalters und der
(2010). Studi mikroanatomi pankreas Prasetyaningtyas, W. E., et al. (2007). Steinzeit 47/48: 29–50.
babirusa (Babyroussa babyroussa) A lectin histochemical study on the Berger, E. M., Leus, K. Vercammen, P.
menggunakan metode pewarnaan testis of the Babirusa, Babyrusa baby- & Schwarzenberger, F. (2006). Faecal
baku dan immunohistokimia. Jurnal roussa (Suidae), Anatomia, Histologia, steroid metabolites for non-invasive
Kedokteran Hewan 4(2): 49–52. Embryologia 36: 343–348. assessment of reproduction in
67

Published online by Cambridge University Press

 Part II: Species Accounts

common warthogs (Phacochoerus Guinea. London: John Murray, Vol 2, of Babyrousa babyrussa (Suidae,
­africanus), red river hogs pp. 190–191, 200–205. Mammalia). PhD thesis, The
(Potamochoerus porcus) and babirusa Hamerton, A. E. (1931). Report on the University of Edinburgh.
(Babyrousa babyrussa). Animal deaths occurring in the society’s gar-   (1996). The habitat and diet of the
Reproduction Science 91: 155–171. dens during the year 1930. Proceedings Sulawesi babirusa (Babyrousa baby-
Blouch, R. (1990). Report from the field: of the Zoological Society of London russa). In Manansang, J., Macdonald,
Indonesia. Conservation and Research 1931: 527–555. A. A., Siswomartono, D., Miller, P. & Seal,
Center Newsletter. Washington, DC:   (1941). Report on the deaths occurring U.(eds.), Population and habitat viability
Smithsonian Institution, 1: 6–8. in the society’s gardens during the assessment for the babirusa (Babyrousa
Bowles, D. (1986). Social behaviour and years 1939–1940. Proceedings of the babyrussa). Report. Apple Valley, MN:
breeding of Babirusa (Babyrousa Zoological Society of London series B Conservation Breeding Specialist Group
babyrussa) at the Jersey Widlife 3: 151–185. (SSC/IUCN), pp. 121–143.
Preservation Trust. Dodo, Journal of Heekeren, H. R. van. (1952). Rock-   (2000). Feeding babirusa (Babyrousa
the Jersey Wildlife Preservation Trust paintings and other prehistoric discov- babyrussa) in captivity. In Nijboer, J.,
23: 86–94. eries near Maros (South West Celebes). Hatt, J.M., Kaumans, W. & Gansloßer,
Burton, J.A. & Macdonald, A. A. (2009). Laporan Tahunan Dinas Purbakala U.(eds.), Zoo animal nutrition. Fürth:
Variation in distribution and abun- 1950: 22–35. Filander, pp. 237–250.
dance of large mammals in Lore Lindu   (1972). The Stone Age of Indonesia. Leus, K. & Macdonald, A. A. (1997). From
National Park, Central Sulawesi, Verhandelingen van het Koninklijk babirusa to domestic pig: the nutrition
Indonesia. Malay Nature Journal 61: Instituut voor Taal-, Land- en of swine. Proceedings of the Nutrition
295–305. Volkenkunde 61: 1–311. Society 56: 1001–1012.
Chaudhuri, M., Carrasco, E., Kalk, P. & Hooijer, D. A. (1948). Pleistocene verte- Leus, K. & Vercammen, P. (1996).
Thau, R. B. (1990). Urinary oestrogen brates from Celebes; III. Anoa depressi- Behaviour of a male and female babi-
excretion during oestrus and preg- cornis (Smith) subsp., and Babyroussa rusa (Babyrousa babyrussa, Suidae,
nancy in the babirusa. International babirussa beruensis nov. subsp. Mammalia) during the first five days
Zoo Yearbook 29: 188–192. Koninklijke Nederlandische Akademie after their move to a semi-natural
Clauss, M., Nijboer, J., Loermans, J. H. M., van Wetenschappen 51: 1322–1330. enclosure. Der Zoologische Garten,
et al. (2008). Comparative digestion   (1950). Man and other mammals from N.F. 66(3): 133–155.
studies in wild pigs at Rotterdam Zoo. Toalian sites in south-western Celebes. Leus, K., Bowles, D., Bell, J. & Macdonald,
Zoo Biology 27: 305–319. Verhandelingen der Koninklijke A. A. (1992). Behaviour of the babirusa
Clayton, L. & MacDonald, D. W. (1999). Nederlandische Akademie van (Babyrousa babyrussa) with ­suggestions
Social organization of the babirusa Wetenschappen 46(2): 1–164. for husbandry. Acta Zoologica et
(Babyrousa babyrussa) and their use   (1954). Pleistocene vertebrates from Pathologica Antverpiensia 82: 9–27.
of salt licks in Sulawesi, Indonesia. Celebes. VIII. Dentition and skeleton Leus, K., Bland, K. P., Dhondt, A. A. &
Journal of Mammalogy 80: 1147–1157. of Celebochoerus heekereni hooijer. Macdonald, A. A. (1995). Ploughing
Clayton, L. M., Keeling, M. & Milner- Zoologische verhandelingen uitgegeven behaviour of the babirusa (Babyrousa
Gulland, E. J. (1997). Bringing home door het Rijksmuseum van Naturlijke babyrussa) suggests a scent ­marking
the bacon: a spatial model of wild pig Historie te Leiden 24: 1–52. function. Journal of Zoology
hunting in North Sulawesi, Indonesia. Indonesian Ministry of Forestry/IUCN 238: 209–219.
Ecological Applications 7: 642–652. (2013). National Conservation Strategy Leus, K., Goodall, G. P. & Macdonald,
Conklin, N. L., Dierenfeld, E. S. & and Action Plan for Babirusa. [Strategi A. A. (1999). Anatomy and ­histology
Maclaughlin, K. (1994). Digestibility dan rencana aksi konservasi Babirusa of the babirusa (Babyrousa ­babyrussa)
and passage of a zoo diet fed to (Babyrousa babyrussa) tahun 2013– stomach. Comptes Rendus de
Babirusa (Babyrousa babyrussa). 2022. Peraturan Menteri Kehutanan l'Académie des Sciences Série III –
Zoologische Garten 64: 357–365. Republik Indonesia Nomor: P.55/ Sciences de la Vie 322: 1081–1092.
Deninger, K. (1909). Über Babirusa. Menhut-II/2013]. Leus, K., Morgan, C. A. & Dierenfeld,
Berichten der naturforschenden Kneepkens, A. F. L. M. & Macdonald, A. E. S. (2001). Nutrition. In Fischer, M.
Gesellschaft zu Freiburg im Breisgau 17: A. (2010). Vertebral column, rib and (ed.), Babirusa (Babyrousa babyrussa)
179–200. sternal muscles of Sulawesi babirusa husbandry manual. St Louis, MN:
Eriawati, Y. (2003). Lukisan di gua-gua karst (Babyrousa celebensis). Anatomia, American Association of Zoos and
Maros-Pangkep, Sulawesi Selatan: gam- Histologia, Embryologia 40: 149–161. Aquariums, pp. 11–28.
baran penghuni dan matapencahariannya Kneepkens, A. F. L. M., Badoux, D. M. & Leus, K., Macdonald, A. A., Goodall,
: album seni budaya [= Rock paintings Macdonald, A. A. (1989). Descriptive G., Veitch, D., Mitchell, S. &
on Maros-Pangkep karst caves, South and comparative myology of the fore- Bauwens, L. (2004). Light and electron
Sulawesi : a depiction of their inhabit- limb of the babirusa (Babyrousa baby- ­microscopy of the cardiac gland
ants and subsistence : Album of Art and russa L. 1758). Anatomia, Histologia, region of the stomach of the babirusa
Culture]. Department of Preservation Embryologia 18: 349–365. (Babyrousa babyrussa – Suidae,
and Development of Culture Ministry of Lee, R. J., Gorog, A. J., Dwiyahreni, Mammalia). Comptes Rendus Biologies
Cultural Media Development. A., et al. (2005). Wildlife trade and 327: 735–743.
Frankham, R., Ballou, J. D. & Briscoe, D. implications for law enforcement in Leus, K., Macdonald, A., Burton, J., &
A. (2010). Introduction to conserva- Indonesia: a case study from North Rejeki, I. (2016). Babyrousa celebensis.
tion genetics. Cambridge: Cambridge Sulawesi. Biological Conservation The IUCN Red List of Threatened
University Press. 123: 477–488. Species 2016: e.T136446A44142964.
Guillemard, F. H. H. (1886). The cruise of Leus, K. (1994). Foraging behaviour, Lopez Gonzales, M. & Taylor, D. C.
the Marchesa to Kamschatka and New food selection and diet digestion (1998). Cutaneous aspergillosis in
68

Published online by Cambridge University Press

 Chapter 6: Sulawesi babirusa Babyrousa celebensis (Deninger, 1909)

a Babirusa (Babyrousa babyrussa). in babirusa (genus Babyrousa). Reksowardojo, D. H. (1995). Ringkasan

Veterinary Record 143: 568. Journal of Zoo and Aquarium Research disertasi studi kemampuan reproduksi
Macdonald, A. A. (1991). Monographie 4: 22–29. dan produksi babirusa (Babyrousa
des hirschebers (Babyrousa babyrussa). MacKinnon, J. (1981). The structure and babyrussa celebensis Deninger) melalui
Bongo, Berlin 18: 69–84. function of the tusks of the babirusa. upaya budidaya. PhD thesis, Bogor,
  (1993). The babirusa (Babyrousa Mammal Review 11: 37–40. Indonesia.
babyrussa). In Oliver, W. L. R.(ed.), Maclaughlin, K., Ostro, L. E. T., Koontz, Sarasin, P. & Sarasin, F. (1905).
Pigs, peccaries and hippos: status survey C. & Koontz, F. (2000). The ontogeny Materialien zur Naturgeschichte
and conservation action plan. Gland: of nursing in Babyrousa babyrussa and der Insel Celebes. Wiesbaden: C. W.
IUCN, pp. 161–171. a comparison with domestic pigs. Zoo Kreidels, Vol. 5, pp. 4–5, 36–43, 52–54.
  (1994). The placenta and cardiac fora- Biology 19: 253–262. Smiet, F. (1982). Threats to the spice
men ovale of the babirusa (Babyrousa Maklis, T. 1991. Rahasia harta yang islands. Oryx 14: 323–328.
babyrussa). Anatomy and Embryology hilang. Jakarta: Pustaka Utama Grafiti. Thenius, E. (1970). Zur Evolution und
190: 489–494. Melisch, R. (1994). Observation of swim- Verbreitungsgeschicht der Suidae
  (2008). Ökologie und Schutz des ming babirusa Babyrousa babyrussa (Artiodactyla, Mammalia). Zeitschrift
Hirschebers. In Macdonald, A. A. & in Lake Poso, Central Sulawesi, fur Saugetierkunde 35:
Gansloßer, U.(eds.), Wilde Schweine Indonesia. Malayan Nature Journal 321–342.
und Flußpferde. Fürth: Filander Verlag, 47: 431–432. United States Department of the
pp. 137–154. Milner-Gulland, E. J. & Clayton, L. Interior. (1980). Republication of the
Macdonald, A. A. & Kneepkens, A. F. L. M. (2002). The trade in babirusa and lists of endangered and threatened
(1995). Descriptive and comparative wild pigs in North Sulawesi, ­species and correction of technical
myology of the hindlimb of the babi- Indonesia. Ecological Economics errors in final rules. Federal Register
rusa (Babyrousa babyrussa L. 1758). 42: 165–183. 45: 33768–33781.
Anatomia, Histologia, Embryologia 24: Mohr, E. (1960). Wilde Schweine. Neue Urbain, A., Bullier, P. & Nouvel, J. (1938).
197–207. Brehm-Bucherei no. 247. Wittenberg- Au sujet d’une petite épizootie de
Macdonald, A. A. & Leus, K. (1995). Lutherstadt: Ziemsen Verlag. Fièvre aphteuse ayant sévi sur des
Creating a public understanding of Munro, S. A., Kaspe, L., Sasmita, animaux sauvages en captivité. Bulletin
the biology of the babirusa (Babyrousa R. & Macdonald, A. A. (1990). de l’Académie Vétérinaire de France
babyrussa) within a caring zoo envi- Gastrointestinal helminthosis in the 11: 59–73.
ronment. Ibex Journal of Mountain babirusa (Babyrousa babyrussa) and Van den Bergh, G. D., de Vos, J. & Sondaar,
Ecology 3: 37–40. response to ‘Albendazole’. Veterinary P. Y. (2001). The late ­quaternary
Macdonald, A. A., Bell, J., Munro, S. A., Record 126: 16. palaeogeography of mammal evolution
et al. (1989). Observations on the Ogle, S. (2010). Techniques to assist in the Indonesian archipelago.
behaviour and health of captive babi- conservation breeding of the babirusa Palaeogeography, Palaeoclimatology,
rusa. In Prociding Simposium Nasional (Babyrousa babyrussa). PhD thesis, Palaeoecology 171: 385–408.
Penyakit Satwa Liar. Surabaya, The University of Edinburgh. Van Wees, S. Nujboer, J. Everts, H. &
Indonesia: Universitas Airlangga dan Ogle, S. & Macdonald, A. A. (2008). Beynen, A. C. (2000). Apparent
Kebun Binatang, pp. 244–253. Die Fortpflanzungsbiologie im digestibility of macronutrients
Macdonald, A. A., Bowles, D., Bell, J. & Überblick [Biology of reproduction in captive babirusa (Babyrousa
Leus, K. (1993). Agonistic behaviour in wild pigs, peccaries and hippos: babyrussa). Journal of Zoo and Wildlife
in captive babirusa (Babyrousa baby- a review]. In Macdonald, A. A. & Medicine 31: 508–511.
russa). Zeitschrift für Säugetierkunde Gansloßer, U. (eds.), Wilde Schweine Vercammen, P. (1991). Ubersicht uber die
58: 18–30. und Flußpferde. Fürth: Filander Verlag, haltung von hirschebern (Babirousa
Macdonald, A. A., Leus, K., Florence, A., pp. 33–66. babyrussa celebensis L.) im Zoo
Clark, J. & Patry, M. (1996). Notes on Patry, M. (1990). Babiroussa: une Vie Antwerpen. Bongo, Berlin 18: 244–249.
the behaviour of Sulawesi warty pigs Jusqu'an Bout du Reve. Paris: Fixot. Whitmore, T. C., Tantra, I. G. M. &
(Sus celebensis) in North Sulawesi, Patry, M., Leus, K. & Macdonald, A. A. Sutisna, U. (1989). Tree flora of
Indonesia. Malayan Nature Journal 50: (1995). Group structure and Indonesia: check list for Sulawesi.
47–53. behaviour of babirusa (Babyrousa Bogor: Forest Research and
Macdonald, A. A., Burton, J. & Leus, K. babyrussa) in Northern Sulawesi. Development Centre.
(2008a). Babyrousa celebensis. The Australian Journal of Zoology Whitten, A. J., Mustafa, M. & Henderson,
IUCN Red List of Threatened Species 43: 643–655. G. S. (2002). The ecology of Sulawesi.
2008: e.T136446A4292795. http:// Perry, G. (1811). Arcana; or the museum Hong Kong: Periplus Editions.
dx.doi.org/10.2305/IUCN.UK.2008. of natural history . . . Printed by Ziehmer, B., Ogle, S., Signorella, A.,
RLTS.T136446A4292795.en George Smeeton for James Stratford, Knorr, C. & Macdonald, A. A. (2010).
.Downloaded 15 December 2015. London. Anatomy and histology of the repro-
Macdonald, A. A., Mitchell, S., Signorella, Peters, C. T. M. (1985). De babiroesa: het ductive tract of the female Babirusa
A., & Leus, K. (2008b). Ultrastructural hertzwijn. Dieren 5: 135–139. (Babyrousa celebensis). Theriogenology
characterization of the epithelium that Purchas, S. (1625). Hakluytus posthumus 74: 184–193.
constitutes the cardiac gland epithe- or Purchas his pilgrimes. London: Ziehmer, B., Signorella, A., Kneepkens,
lium ‘honeycomb’ in the stomach of Henry Fetherstone. A. F. L. M., et al. (2013). The anatomy
the babirusa (Babyrousa babyrussa). Reinhard, R. & Frädrich, H. (1983). and histology of the reproductive tract
Comptes Rendus Biologies 331: 32–41. Bemerkungen zur Zucht Hirschebern of the male Babirusa (Babyrousa
Macdonald, A. A., Leus, K. & Hoare, H. (Babyrousa babyrussa L.) im Zoo celebensis). Theriogenology 79:
(2016). Maxillary canine tooth growth Berlin. Bongo, Berlin 7: 65–70. 1054–1064.
69

Published online by Cambridge University Press