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Pigs, Peccaries and
Hippos Status Survey and Action Plan (1993) Chapter 5 The Eurasian Suids Sus and Babyrousa 5.1. Taxonomy and Description Colin P. Groves and Peter Grubb The Genus Sus According to the major review of the genus Sus by Groves (1981) at least 5
species and 24 subspecies are recognized. A more recent study by
Hardjasasmita (1987), is at variance with Groves' review, in that it
recognizes several additional species and subspecies from Indonesia. However,
as this author does not provide any new evidence to support his conclusions
or to counter those of Groves' we do not see any need to modify the latter's
treatment of the wild pigs of this country. Nevertheless, some important new
evidence suggests that two additional species should be recognized from the
Philippines (see below), making a total of 7 species and (approximately) 22
subspecies, as follows: 1.
The Eurasian
Wild Pig, Sus scrofa (c. 17 ssp.). 2.
The Pigmy Hog, Sus salvanius (0 ssp.). 3.
The Javan Warty
Pig, Sus verrucosus (2 ssp.). 4.
The Bearded
Pig, Sus barbatus (3 ssp.). 5.
The Philippine
Warty Pig, Sus philippensis (0?
ssp.). 6.
The Visayan
Warty Pig, Sus cebifrons (0? ssp.). 7.
The Sulawesi
Warty Pig, Sus celebensis (0 ssp.). Groves (1981) also drew attention to a 'mystery'
pig, S. bucculentus, which is known
from only two skulls from Cochin China (now extreme south Viet Nam),
collected in the late 19th Century. This animal is evidently closely allied
to S. verrucosus, but its external
appearance and present status are entirely unknown. The pigs of this genus may be separated into two
groups, the non-warty pigs (2 spp.) and the warty pigs (5 or 6 spp.), these
groups being differentiated by various cranial and dental characters and the
development of conspicuous facial adornments or 'warts' in adult males of the
warty pigs alone. The Non-warty Pigs of the Genus Sus The two species are sympatric, S. scrofa occurring over the entire known range of S. salvanius, namely the sub-Himalayan
alluvial tract of the northern Indian sub-continent, so they must be
specifically distinct. Moreover, S.
salvanius has a number of highly distinctive characters (e.g. extreme
reduction in body size, rudimentary tail and only three pairs of mammae),
which serve to differentiate it from scrofa
and, indeed, all other wild pigs. These characters have led some authors to
annex the species as a separate genus or sub-genus, Porcula, though Groves considered it more closely allied to scrofa than to other members of the
genus, and believed that most of the species' diagnostic features are
correlated with the reduction in body size. In both species the male's lower canine has the
inferior surface narrower than the posterior surface. The female's canines
are like small versions of the male's. The preorbital fossa is shallow with
sloping borders. 1. The Eurasian Wild Pig (Sus scrofa) A moderate to large sized pig with a relatively
short muzzle and no face warts. Females 88.8-95.3% of males in condylobasal
length in various populations; about 80% of male body weight. Relatively
long-limbed, especially in northern races; medial false hooves as long as
lateral; ears relatively large; tail relatively long, with simple tuft; head
not large compared to body; snout disc perpendicular to axis of head; back
rounded. No warts or gonial whorl. Often well-haired; hairs all agouti;
general color brown with a tendency to blackish, grayish or rufous; face,
cheeks and throat with a grizzling of whitish hairs or more strongly
expressed markings; underside dark; bristles of nape long, in some races
forming a distinct narrow nuchal crest or mane which extends down the back;
under-fur thick and woolly or absent. Piglets longitudinally striped, stripes
gradually fading after five to six months. Four to six pairs of mammae. There is considerable variation in size, with
sample-means for greatest length of skull in males ranging from 275 mm
(Ryukyu Islands) to 466 mm (Ukraine). In general, insular pigs are smaller
than those on the adjacent mainland, and southern pigs are smaller than
northern ones. In Groves's (1981) revision of this species, no groupings of
subspecies were recognized, but four regional groupings are used here in an
informal sense in order to distinguish certain features amongst the 17
currently recognized subspecies (Fig. 10, section 5.2): ·
Western races.
Mane poorly developed except in the nominate race and adjacent attila; under
wool thick. Sample-means of greatest length of skull in males 308-466 mm.
Mostly high-skulled races, though lybicus and some scrofa are low skulled. The
following (6) subspecies are included: S. s. scrofa (northern low-skulled and southern high-skulled populations may be
subspecifically distinct): North Spain, North Italy, France, Germany,
Benelux, Denmark, Poland, Czechoslovakia, Albania (?); S. s. meridionalis: Andalusia; also Sardinia & Corsica (though the island
populations are almost certainly feral, having arrived in the Holocene,
probably as primitive domestic stock - see Vigne, 1988); S. s. algira: Tunisia, Algeria, Morocco on the coastal side of the mountains; S. s. attila: Hungary, Ukraine, Central Byelorussia east into Soviet Central Asia
as far as the Aral Sea; south to the northern flank of the Caucasus and the
Mesopotamian Delta, Iraq; S. s. lybicus: Transcaucasia, Turkey, Levant, Palestine, Yugoslavia (also extinct
in Nile Delta, though this was probably a feral population – see: Verpmann,
1987 - as there is no record of this species of the species in Egypt in any
archeological sites of Noelithic, Bronze or Iron Age); and S. s. nigripes: flanks of the Tienshan Range in Central Asia and (according to
Russian authorities) ranging west to the Caspian Sea, south to northern Iran,
Afghanistan, western and southern Mongolia and China, and east as far as
Novosibirsk. ·
Indian races.
Under wool sparse (davidi) or
absent; mane long and thick; snout and mouth-gonion bands fairly well defined
(davidi) or absent. Sample-means of
greatest length of skull in males 365-490 mm (339 mm in Bopeta - see below).
All are high-skulled races, except for davidi, which is low skulled. The (4)
subspecies included are: S. s. davidi: Pakistan and north-west India to south-eastern Iran; S. s. cristatus: Sub-Himalayan region from Ludhiana, Punjab, through Nepal and Sikkim
to Nagaland and North Burma; Uttar Pradesh; Madhya Pradesh; probably
Kolhapur; Bihar; Bengal south through Burma and western Thailand to Isthmus
of Kra; S. s. affinis: southern India and most of Sri Lanka; and S. s. subsp: Bopeta, Central Province, Sri Lanka (based on a single very small
skull). ·
Eastern races.
A definite whitish streak from angle of mouth to lower jaw; under wool thick
except in moupinensis; mane poor or absent. Sample-means of greatest length
of skull in males 327-474 mm (275 on Iriomote). Mostly high-skulled races,
except for ussuricus. The (6) subspecies included are: S. s. sibiricus: Mongolia, Transbaikalia; S. s. ussuricus: Soviet Far East, Manchuria, Korea; S. s. leucomystax: Honshu, Shikoku, Kyushu; S. s. riukiuanus: Ryukyu Islands (including at least Ishigaki and Iriomote; S. s. taivanus: Taiwan; S. s. moupinensis: Coastal China south to Vietnam and west to Szechwan. ·
Indonesian race
(or banded pig), S. s. vittatus.
Body hair sparse, no under wool, mane fairly long, a broad reddish brown band
along middle of muzzle, broadening at angles of mouth and on sides of upper
lip, where it extends backwards to disappear on sides of neck Greatest length
of skull in males, 284-380 mm in sample means. Range: Peninsular Malaysia;
the offshore islands of Terutau and Langkawi; Sumatra; Riau Archipelago
(includes Bulan, Jeri, Ungar Batam and Karimun); Java and offshore island
Peucang; Bali; Lombok; Sumbawa; and Komodo. 2. The Pigmy Hog (Sus salvanius) This is the smallest of the living suids, with
greatest length of skull in males averaging only 179 mm; body weight is
6.6-9.7 kg in adults, head and body length 55-71 cm (mean, 58 cm) (Oliver
1980); female about 87% of male in condylobasal length. Relatively
short-limbed; hind limbs long relative to forelimbs; medial false hooves
relatively short; ears small; tail very short, barely tufted; head moderate
in size; snout disc perpendicular to axis of head; back short, rounded. No
warts or gonial whorl. Hairs all agouti-banded; little or no mane; underside
wholly dark. Piglets faintly striped. Only three pairs of mammae. The pigmy
hog is confined to the tall grass savannah or terai of the Himalayan
foothills and now restricted to northern Assam (Oliver, 1980). The Warty Pigs of the Genus Sus The warty pigs have sometimes been referred to as
the 'verrucosus group' distinct from the two, non-warty species (above), one
of which (scrofa) is sympatric with verrucosus and barbatus over parts of
their ranges. Hybridization between scrofa and all five species of warty pigs
has been recorded, albeit in captivity in most instances. Since all of the
warty pigs have parapatric distributions, they are considered conspecific by
some authors and, unlike the non-warty Sus spp., their separation is based
exclusively on morphological characters. The 'warty pigs' are so-called because the adult
males typically develop three pairs of warts, namely 'infraocular' (=
infraorbital) warts on the cheek (malar) swellings; 'gonial' warts on the jaw
angle; and 'preocular' (= preorbital) warts above the canine root flanges.
The relative development of these warts, which continue to enlarge throughout
life, are diagnostic for three 'Sundaic' species, the infraorbital warts
being most developed in verrucosus and the preocular warts being most
developed in celebensis. The facial warts are reduced in barbatus, in which
the gonial warts are concealed by the growth of the cheek whiskers, the
'gonial whorl' or 'beard', and in the two eastern (Wallacian) Philippine
species, philippensis and cebifrons. The warts may also be
present, though very small, in females of these species. The structure of the
canine is also characteristic of warty pigs, the inferior surface of the
lower canine being broader than the posterior surface, so that a section of
the tooth is more or less an equilateral triangle. The females have
diminutive canines, which tend to point simply down (upper teeth) or up
(lower). The preorbital fossa is always deeply excavated with over-hanging,
shelf-like borders. 3. The Javan Warty Pig (S. verrucosus) This species and the next are distinguished by the
great elongation of the face. Size medium to large; sexual dimorphism in size
greater than in other species; males very large - more than twice the weight
of females; female condylobasal skull length 80.3 and 83.3% of male
measurements in two populations; relatively long limbed; ears large; tail
relatively long, tufted simply; head large, heavy compared to body; back
long, straight; snout disc somewhat oblique to axis of head. Three pairs of
warts; infraorbital pair largest; long tuft of hair on gonion marks spot
where gonial wart will emerge late in life in males. Body hairs red to yellow
with black tips, or pure black; a broad mane extending to rump; underside
sharply marked off, yellowish. Piglets faintly striped, when very small only.
Six pairs of mammae. Two subspecies are recognized, as follows: S. v. verrucosus: Java and, formerly, Madura,
where it is now extinct; and S. v. blouchi: confined to Bawean The nominate form is much larger, with sample-means
of greatest skull length 408-429 mm in males; the recently described Bawean
race (Groves, 1981) having a skull length of only 354 mm. 4. The Bearded Pig (S. barbatus) S. barbatus is closely related to S. verrucosus, but is typically
distinguished by the bushy gonial tuft of hair enlarged into a cheek-beard;
much smaller warts; and often a still more elongated skull. Size medium to
large; females relatively larger in relation to males, 88.2-93.6% of males in
condylobasal skull length. Relatively long limbed; ears relatively small;
tail long with large terminal tuft divided into distinct anterior and
posterior parts; head large, heavy; snout disc quite oblique to axis of head;
back straight, long. Two pairs of warts in males; whorl on gonion but no gonial
wart; warts small or absent in females. Hairs agouti or uniform (black);
broad long mane, extending to rump; underside wholly dark. Piglets striped,
but pattern fading before six months. Six pairs of mammae. The three
currently recognized subspecies, are: oi: Peninsular
(West) Malaysia, Sumatra, Bangka and Palau Bintang in the Riau Archipelago; barbatus: Borneo and the westernmost of the Sulu Archipelago (Sibutu and
Tawitawi); and ahoenobarbus from Balabac, Palawan and offshore islands, and the Calamianes group. The subspecies differ in size, coloration and hair
development. The nominate race of Borneo is large, with a well-developed
beard. S. b. oi is somewhat
variable over its disjunct range, but is also very large, has coarse bushy
hair over the top of the snout and a smaller beard. S. b. ahoenobarbus is much smaller and darker. Sample-means of
greatest length of skull in males are 359-365 mm, against 435-505 mm in the
larger races. 5. The Philippine Warty Pig (S. philippensis) Following Sanborn (1952), the wild pigs of the main
(Wallacian Region) Philippine archipelago (i.e. excluding the Palawan group
of islands in the Sundaic Region) have generally been assigned to the species
S. celebensis. However, Groves
(1981) rejected this arrangement on finding that their cranial characters
aligned them with S. barbatus and
not S. celebensis - a view
subsequently endorsed by Mudar (1986) - whilst acknowledging that they might
merit separation as full species. Recently-photographed captive adult boars from
Panay (S. 'barbatus' cebifrons) and
from Luzon and Mindanao (S. 'barbatus'
philippensis) are unlike S.
barbatus and differ amongst themselves. Furthermore, skulls of S. 'b.' cebifrons from Negros in the
central Philippine 'Visayas Region' recently examined by C. P. Groves show
several differences from skulls of philippensis.
This new information is yet to be fully reported but it has led to a
reappraisal of the systematics of Philippine pigs. It now seems appropriate
to recognize two separate species on the Philippines (excluding the Palawan
Region), namely S. philippensis and
S. cebifrons - a treatment that
corresponds with the systematics of the Philippine deer (Grubb and Groves,
1983). The Philippine warty pig, S. philippensis, occurs on most of the larger, outermost islands
(Mindoro, Mainit, Luzon, Catanduanes, Samar, Leyte, Biliran, Mindanao,
Basilan and Jolo) in the eastern Philippines. It is a black pig, sometimes
with a pale snout-band and red-brown patches in the mane. It is smaller than S. barbatus ahoenobarbus but somewhat
larger than S. cebifrons (means of
greatest skull-length measurements are 318-335 mm in adult males). A boar
from Luzon strongly resembles S.
celebensis in the distinct crest and prominent preorbital warts, though
these features are less strongly expressed in another male from Mindanao.
This variation may reflect differences between island populations, a
conclusion that is to be expected in view of faunal provincialism in the
Philippines (Heaney, 1986). Therefore, it will be necessary to reconsider the
systematics of the species, as it may prove to be polytypic. 6. The Visayan Warty Pig (S. cebifrons) This is a smaller pig occurring allopatrically to S. philippensis on the west central
islands of the Philippines. Skull length is 299 mm (mean of two males from
Negros) and recently determined features of Negros skulls distinguish this
species from S. philippensis as
well as from other, better-known warty pigs. The external appearance is not
well known but boars from Panay have a long mane extending to the rump and
large facial warts, while skins from Negros have white hairs on the shoulders
and sides. The species is reliably recorded from the Visayan Islands of Cebu,
Siquijor, Guimaras, Negros and Panay - identified as the Negros Faunal Region
by Heaney (1986). However, it is already extinct over the majority of its
known, former range (see later text). 7. The Sulawesi Warty Pig (S. celebensis) This is a short-faced species, like the Eurasian
wild pig, of small size (sample-means of greatest length of skull 256-331 mm
in males, females 85.4-90.4% of males in condylobasal skull length in
different populations), though there is some evidence of a north (smallest)
to south (largest) cline in body size on mainland Sulawesi, extending onto
the offshore islands. The species is characterized by its (relatively) very
short legs; small, short ears; long, simply tufted, tail; large, heavy head;
short, slightly convex, back; and somewhat oblique snout disk. The preorbital
wart is typically the most developed of the three pairs, though the gonial
wart, which is marked only by a whorl in younger animals, does become obvious
late in life and may hypertrophy. The warts remain small in females. The
pelage is usually black, often with white or yellow hairs intermixed, but
some specimens are predominantly red-brown or even yellowish in color, the
underside being uniformly dark in young animals, but becoming paler,
eventually light yellow, with age. A distinct yellow snout band is usually
present. The mane is rather poorly developed, but forms a conspicuous tuft or
'crest' on the crown, especially in young adult males. Piglets striped to at
least six months of age. There are six pairs of mammae. The distribution of this species is most unusual (Fig.15,
section 5.7). It occurs as a native form only on Sulawesi and offshore
islands (i.e. the Togian Islands, Pulau Salayer, P. Butung, P. Peleng and P.
Lembeh), but it has been introduced onto Buru, Halmahera and Seram, and some
of the Lesser Sunda Islands (Timor, Flores and P. Roti) and the west Sumatran
Islands (Simaleue and Nias) as a domesticated or feral form. Indeed, as
Groves has shown, it is evident that the species has been truly domesticated
(probably in the early Holocene on Sulawesi), and transported to the
Moluccas, where it has also hybridized with S. scrofa, thereby giving rise to the second species of Sus ever
to be described, 'S. papuensis', the domestic and feral pig of New Guinea.
The so-called 'S. timorensis' (from
Timor), 'S. floresianus' (Flores), 'S. mimus' (Simeulue) and 'S. niadensis' (Nias) are also all
variously modified celebensis-pigs,
some of which are still maintained as domesticates, whilst others (like those
on Buru and Seram) were either released to be hunted whenever required for
eating or have been allowed to revert to the wild state. The Genus Babyrousa 8. The Babirusa (B.
babyrussa) This extraordinary animal is confined to Sulawesi
and some neighboring islands. Three extant subspecies are widely recognized
(Groves, 1980). These are: B. b. babyrussa: Buru and the Sula Islands
(Mangole, Taliabu and, formerly, Sulabesi). B. b. togeanensis: Togean Islands (Malenge, Talatakoh, Pulau
Batudaka, Batone and Pangempong). B. b. celebensis: northern peninsula of Sulawesi
at least as far west as Bumbulan and from Pulau Lembeh. A fourth subspecies, B. b. bolabatuensis, described from an early Holocene (?) skull
from Bola Batu caves near Watampone, may also survive in still-forested areas
of central and south Sulawesi. Indeed, a recent skull from Gunung Melema,
Moa, near Kulawi, could be ascribed to this race, though it also resembles
the skulls of the Buru and Sula Islands' animals, which may have originated
from this region of Sulawesi (see later text). The subspecies differ in size, hairiness and
characters of the skull and teeth. Leaving aside the problematical bolabatuensis, nominate babyrussa is the smallest and the
hairiest. Indeed, this subspecies is sometimes referred to as the 'golden' or
'hairy' babirusa, on account of its relatively long (>5cm), yellowish
pelage, in marked contrast with the barely discernable, short, sparse pelage
of the most familiar form, B. b. celebensis, from the (north) Sulawesi
mainland. The latter subspecies is also the largest, the Togian Islands', togeanensis, being intermediate in
body size between these forms. References Groves, C.P.
1980. Notes on the systematics of Babyrousa
(Artiodactyla, Suidae), Zool. Meded. Leiden, 55: 29-46. Groves, C.P.
1981. Ancestors for the Pigs: Taxonomy and Phylogeny of the Genus Sus. Tech. Bull., No. 3, Dept. of
Prehist., Research School of Pacific Studies, Australian National University:
96pp. Grubb, P. and
Groves, C. P. 1983. Notes on the taxonomy of the deer (Mammalia, Cervidae) of
the Philippines. Zool. Anz., Jena 219 (1/2): 119-144. Hardjasamita,
H. S. 1987. Taxonomy and phylogeny of the Suidae (Mammalia) in Indonesia.
Scripta Geol., 85: 1-68 Heaney, L. R.
1986. Biogeography of mammals of S. E. Asia: estimates of rates of
colonisation, extinction and speciation. Biol. J. Linnean Soc. 28: 127-165. Mudar, K. M.
1986. A morphometric analysis of the five subspecies of Sus barbatus, the bearded pig. (Unpubl.) M. Sc. thesis, Michigan
State University. Oliver, W.L.R.
1980. The Pigmy Hog: the Biology and Conservation of the Pigmy Hog, Sus (Porcula) salvanius, and the
Hispid Hare, Caprolagus hispidus. Spec. Scien. Rep. No. 1, Wildlife
Preservation Trust, Jersey,: 80pp. Sanborn, C. C.
1952. Philippine Zoological Expedition 1946-47. Fieldiana: Zoology, 33: 89-158. Verpmann, H. P.
1987. The ancient distribution of ungulate mammals in the Middle East.
Beihefte zum Tübinger Atlas des vorderen Orients, Reihe A
(Naturwissenschaften), 27: 1-173. Vigne, J.- D. 1988. Les mammifères post-glaciaires
de Corse. Gallia Prehistoire, 26th supp.: 334 pp. |
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Eurasian Wild Pig |