Pigs, Peccaries and Hippos Status Survey and Action Plan (1993)

 

Chapter 4.4

 

The Bush Pigs

(Potamochoerus porcus and P. larvatus

 

Paul Vercammen, Armin H. W. Seydack and William L. R. Oliver.

 

 

Status and Action Plan Summary

 

Status categories 1-2 (widespread and locally abundant or relatively secure) - both species.

 

The bushpig, Potamochoerus larvatus, and red river hog, P. porcus, remain relatively widely distributed and are locally abundant in some places, and neither is considered threatened over the majority of their known ranges at the present time. Both species live in small family groups, usually comprising 4-10 individuals, and are rarely observed in larger numbers. They are usually sedentary and territorial. In most areas they are predominately nocturnal, but tend towards diurnalism if undisturbed. Reproduction is seasonal with litter size from one to six, but generally only one or two young are reared successfully. Both species, but particularly bushpigs, are notorious for their depredations on crops. There is evidence that the conversion of former forest to secondary scrub and agriculture has resulted in an increase in their numbers in some areas, and attempts to control or eradicate them have usually proved unsuccessful.

 

As neither species is considered threatened, priority recommendations are primarily directed towards: a) the resolution of outstanding questions pertaining to their systematic relationships and the distinctness of some regional populations of P. larvatus; b) the need to conduct more field studies on their ecology, behaviour, management and socio-economic importance to local people; and c) obtain additional field status data in areas where they remain poorly known.

 

 

Introduction

 

Following Grubb (this vol. section 4.1), two species of 'bush pigs' are recognized: the red river hog Potamochoerus porcus (0 ssp.) and the bushpig, P. larvatus (4 + ? ssp.), though the genetic and distributional relationships of the two species remain uncertain in the interface between their respective known ranges.

 

Both species are of medium body size, with an elongated snout and a long, often brightly colored coat. The red river hog, P. porcus, is always bright rufous in color, with a distinct white dorsal stripe and crest, long white whiskers and eartufts. The species occurs only in Equatorial [West] Africa, from Senegal in the extreme west, and east and south to eastern Zaire (now Democratic Republic of Congo). Geographical variation is slight, though there is an east (largest) to west (smallest) cline in body size in specimens from eastern Zaire and Cameroon, respectively. However, any such geographic variation is considered insufficient to warrant subspecific separation (P. Grubb, this vol. and in litt.)

 

The bushpig is light red to brown, gray or predominately black in color, sometimes with, but often without, the distinct white masks and long eartufts characteristic of P. porcus (for a full description of regional variants see Grubb, this vol.). P. larvatus has a relatively wide range, extending from Somalia in the north-east, to east and south Zaire in the west, to Natal and Cape Provinces of South Africa in the south (Fig. 9). The four, currently recognized subspecies are:

 

P. l. hassama, eastern Africa;

P. l. koiropotamus, Angola and south-eastern Africa;

P. l. larvatus from Mayotte (Comoro Is. and western Madagascar); and

P. l. hova from eastern Madagascar.

 

However, the Madagascan/Comoron populations are almost certainly introduced, and there is not yet enough evidence to justify the recognition of two possibly distinct populations in eastern Somalia/northern Kenya (i.e. P. l. 'somaliensis' - herein considered as P. l. hassama) and southernmost, apparently isolated population in Cape Province (i.e. P. l. 'nyasae' - herein considered as P. l. koiropotamus; P. Grubb, pers. comm.).

 

 

Link to Fig. 9: Approximate former and present known distribution of the red river hog, Potamochoerus porcus, and the bushpig, P. larvatus ssp.

 

Former and Present Distribution

 

P. porcus remains relatively widely, but now patchily, distributed through much of it its former range in Equatorial West Africa from Senegal/Gambia in the west, through the Guineo-Congolian rain forest zone, extending northwards into the Sudanian transitional zone (Stuart and Adams, 1990). In central and eastern Africa, it occurs at least as far north as southern Chad and at least as far east as Virunga and Garamba National Parks in eastern Zaire (now Democratic Republic of Congo). The species may also occur in extreme south-western Sudan (Hillman, 1982; J. C. Hillman, pers. comm.), though the animals reported from this country so far are P. l. hassama (P. Grubb, pers. comm.). Further east and in south-eastern Zaire the species is replaced by P. larvatus (see below). However, the precise limits of the ranges (or intergradation ?) of these two species in this region and elsewhere in southern Zaire, and hence also their ecological and systematic relationships, has yet to be established (P. Grubb, pers. comm. and this vol., section 4.1).

 

P. larvatus hassama occurs in Eritrea and Tigre Provinces of Ethiopia in the north, (presumably) extending through south-western Sudan on both sides of the Nile. The subspecies P. l. somaliensis (if valid) replaces hassama in the vicinities of the Tana, Juba and Scebeli Rivers in Somalia, N. E. Kenya, and perhaps as far south as Tendaguru on the coast of north Tanzania. The identity of the bushpigs in central Tanzania remains uncertain, so the northern limits of the range of P. l. koiropotamus are provisionally given as southern Tanzania and south-eastern Zaire. This race has by far the largest range of any of the currently recognized subspecies, though it should be reassigned to 'P. l. nyasae' if the (now ?) isolated population in the southern Cape (the origin of the holotype) is shown to be subspecifically distinct (Grubb, this vol.). It otherwise occurs throughout south-eastern Africa, at least as far south as Natal Province in South Africa, as well as in a few isolated locations in west and central Angola.

 

As previously stated, the origins and distinctness of the Madagascan subspecies, P. l. hova and P. l. larvatus, is problematic. The eastern 'race', P. l. hova, closely resembles P. l. koiropotamus, but is smaller in size than the mainland form and the nominate P. l. larvatus from western Madagascar and Mayotte. Clearly, more specimens are needed to elucidate the validity of these subspecies, which are reported to occur in all terrestrial habitats on Madagascar, except in the environs of the major townships and the deforested central plateau (M. Nicoll and F. Rakotondraparany, pers. comm.).

 

Data on the former distributions of both of these species are sketchy and imprecise. However, there has been an evident contraction in the recent range of P. porcus, particularly in the west and extreme north of its range. The expansion of the Sahel zone and, hence, a reduction in cover and the availability of open water, has resulted in a similar contraction in the range of P. larvatus in parts of N. E. Africa. Nonetheless, both species seem to have maintained their presence over the majority of their former ranges, and recent, localized expansion in their ranges has been reported in some areas, e.g. in Botswana (Lloyd and Millar, 1983).

 

 

Habitat, Ecology and Behavior

 

The distribution of both species is apparently limited by the continuous availability of food, water and cover, and they are only rarely reported in open woodland, savannah or other more arid and open habitats.

 

The only field study to have been conducted to date on P. porcus was by Oduro (1989) in Nigeria. From this and other (mostly anecdotal) sources, it appears that this species occurs in a wide range of habitats, including lowland rain forest, gallery forest, dry forest, savannah woodland, and mixed scrub and cultivated areas.

 

From what is known, P. larvatus also occurs in an astonishingly wide range of habitats from sea-level to montane forest (up to 4,000 m on Mt. Kilimanjaro), to gallery forest, flooded forest and swampland, woodland, and mixed scrub and cultivation areas. The species was the subject of a recent, intensive study by Seydack (1983, 1990, 1991). Earlier accounts of its natural history include those of Phillips (1926), Maberly (1967), Attwell and Bearder (1976) and Breytenbach (1979). General accounts of the species' biology are also provided by Sowls and Phelps (1968), Skinner et al. (1976), Ghiglieri et al. (1982) and Kingdon (1982).

 

Both species are predominately nocturnal, though a clear seasonal trend amongst bushpigs in the southern Cape towards more diurnal activity during winter suggests that temperature regulation, rather than hunting pressure, may be the underlying factor in the daily activity patterns of this species. During periods of inactivity they shelter in dense vegetation, and they may construct bad weather nests during cold and wet spells. Average daily movement distances for bushpigs were found to be 3 km, ranging between 0.5 and 5.8 km (Seydack, 1990).

 

They are omnivorous, and may be potentially important dispersers of seeds. The diet of Cape bushpigs was found to comprise 40% subterranean plant parts (mostly tubers, rhizomes and corms), 30% herbage, 13% fruit, 9% animal matter and 8% fungi (Seydack, 1990). Field studies of this species in Uganda have revealed that bushpigs often associate with groups of foraging monkeys in order to feed on discarded fruits (Ghiglieri et al., 1982). They are also opportunistic predators, and consume a range of invertebrates, smaller vertebrates and carrion.

 

Most reports indicate that both species typically live in family groups of 9 -15 individuals, though P. larvatus group sizes in South Africa were found to range from 1 to 10, with average of 2.4 individuals (Seydack, 1990). Similarly, in Madagascar groups seldom comprise more than 5 individuals (M. Nicoll, pers. comm.). Contrarily, Oduro (1989) recorded P. porcus groups of 1 - 15 (x = 10.56) with an immature/adult age ratio of 2:1 in Nigeria, and much larger groups of 30-60 red river hogs have been observed on occasions in Guinea and E. Zaire (L. Macky and J. Hart, pers. comm.).

 

Population density estimates in the southern Cape ranged from 0.3 to 0.5 bushpigs per sq. km, and average home range sizes in the Knysna Forest ranged between 3.8 and 10.1 sq. km (n = 8), with an average of 7.2 sq. km (Seydack, 1990, 1991). The home ranges of the pair-bonded adults in this study area were spatially exclusive, i.e. resource territoriality was actively maintained through defense and patrolling. This study also confirmed the earlier anecdotal accounts of Maberly (1967), Attwell and Bearder (1976) and Skinner et al. (1976) of a monogamous mating system in which adult boars play an active role in the rearing and defense of the young.

 

Reproduction in both species is apparently seasonal, with piglets being most frequently reported towards the end of the dry season or coinciding with the onset of the rainy season. Nearly 75% of births in the bushpig population in Cape Province studied by Seydack (1990) occurred during the spring (September to November), which agrees with earlier records suggesting farrowing taking place during the spring and summer (Sowls and Phelps, 1968; Tinley, 1977). Available data indicate that both species have a similar gestation period of c. 120 days, and litter size of 1-6, with a mode of 3 (Oduro, 1989; Neurohr, 1991; Seydack, 1991).

 

Mortality due to starvation has been recorded in all age classes of P. larvatus in the southern Cape. However, inclement weather and predation were predominant mortality factors for immature animals according to Seydack's (1990, 1991) findings, whereas intraspecific strife played a significant role in adult mortality.

 

 

Threats to Survival

 

Although habitat destruction is not considered a major threat to the survival of either of these species over most of their ranges at the present time, deforestation coupled with intense hunting pressure has inevitably resulted in marked range contractions in some countries/regions. In Benin, (north) Equatorial Guinea and (possibly) south Sierra Leone, for example, P. porcus is reported to be rare or absent outside existing protected areas (A. Green, pers. comm.; A. Blom, pers. comm.; G. Teleki, pers. comm.), though precise data are difficult to obtain owing to the species' secretive and nocturnal behaviour. Similarly, P. larvatus is reported to be scarce or absent outside the better protected areas in Burundi (P. Chardonnet, pers. comm.), though it is doubtful if this species is as seriously threatened as many of the other larger mammals in this country.

 

Indeed, both species are apparently highly adaptable and may even benefit by the opening up of former forested areas by the creation of secondary habitats, the provision of cultivated foodstuffs and reductions in the numbers of their natural predators. Both bushpigs and river hogs are notorious for their depredations on crops. In Zaire (now Democratic Republic of Congo) and Malawi, for example, bushpigs are reputed to cause more damage to agriculture, particularly maize crops, than any other species (R. Bell, pers. comm.). For these reasons, and because they are sometimes regarded as important vectors of livestock diseases (see below), they are widely persecuted by farmers and they are frequently targeted in wildlife control programmes. Generally speaking, all such eradication attempts have been unsuccessful, largely because of the species' cryptic lifestyle and relatively high reproductive potential.

 

Both species are also hunted widely for subsistence purposes, though they undoubtedly benefit from taboos on the consumption of pork in many parts of their ranges. Amongst some Zambian tribes, for example, bush pig meat is considered to be unhealthy, even dangerous, because the animals are reputed to harbor various diseases, including epilepsy. These beliefs may have been influenced by the spread of Islam, which effectively affords these animals a good deal of protection against hunting in many African countries and in some parts of Madagascar. However, even in predominantly Muslim countries, it is doubtful if they are completely free of hunting pressure, since some Muslim groups discriminate between the 'red' meat of Potamochoerus (which is eaten) and the 'white' meat of other suids (P. Chardonnet, pers. comm.), whilst many others permit hunting by non-Muslim groups (e.g. in Sierra Leone; G. Davies, pers. comm.) or even hunt wild pigs themselves in order to sell the meat (e.g. in Guin‚a Bissau, P. Chardonnet, pers. comm.). Seventy-eight percent of hunters interviewed in Gabon by Lahm (1990) cited the sale of red river hog meat as amongst their most important sources of revenue, and that only about one-third of their gained bush-meat was retained for domestic consumption. In Gabon (M. Nicoll, pers. comm.), Zaire (K. H. Smith, pers. comm.) and Guin‚a Bissau (P. Chardonnet, pers. comm.) most animals killed by hunters and farmers are used for local consumption, though there is some local trade to village or city markets. With the exception of a few live individuals exported to zoos (see later text), there is no international trade in these species, or their meat, hide or other products, as far as is known.

 

Potamochoerus spp. are allegedly host to or vectors of tick-borne diseases, such as trichinosis, African swine fever and probably trypanosomes (W. Odura and F. Amubode, pers. comm.). However, the significance of this has yet to be verified.

 

Dorst and Dandelot (1970), Lever (1985) and Oliver and Brisbin (this vol.) cite records of the occurrence of feral pigs (Sus scrofa) in several parts of Sudan and in South Africa. Some of these animals became naturalized as early as 1925, either as a consequence of allowing domestic stock to range freely or by their deliberate introduction in an effort to control the pine tree emperor moth (Nudaurelia cytheria), whose pupae are readily consumed by these animals (Thomas & Kolbe, 1942). Hybridization between P. larvatus and S. scrofa was recorded in the Transvaal in the early 1970's, when an (escaped) domestic sow was mated by a male bushpig and the resulting progeny (a litter of eight) were reported to have bushpig characteristics (Smithers, 1983). Simoons (1953) also cited references to the interbreeding of Potamochoerus with domestic pigs in the Congo and in the Niger Delta. More recently, Eurasian wild pigs (S. scrofa), introduced for hunting purposes, have become established in a more or less free-ranging state in Burkina Faso, Gabon and, probably, Zaire. In the Wonga-Wongue Presidential Hunting Reserve in Gabon, these animals are reported to have interbred with P. porcus, and that their hybrid offspring are running wild (East, 1990). Whilst it is important to stress that evidence for such intergeneric hybridization remains anecdotal and that putative hybrids have yet to be critically examined and described, introductions of S. scrofa apparently pose a threat to the genetic integrity of local Potamochoerus populations, as well as a risk of disease transmission.

 

 

Conservation Measures Taken

 

Both species are known or are likely to occur in all of the principal protected areas within their respective ranges. In all countries with an existing protected areas network, hunting is forbidden or is restricted to permit holders. In most countries such permits are usually given only for scientific research purposes, although they have also been issued on occasions (e.g. in Malawi; R. Bell, pers. comm.) to reduce the numbers of these animals following repeated complaints from local farmers about their damage to crops in neighboring areas.

 

Outside designated national parks and wildlife reserves, protective legislations are more varied. In Guinea, Liberia and Sierra Leone, for example, Potamochoerus spp. remain unprotected and can be legally hunted at all times, whereas in most other countries they are treated as game animals which can only be hunted during an open season. In only a very few countries, such as Burundi, where they are now confined to a few isolated locations, are they protected at all times and throughout their remaining range. In theory, in those areas where hunting is controlled the number of animals taken by one hunter is limited and/or confined to particular age/sex classes. In Gabon, for instance, hunting permits are issued for no more than 2 adult P. porcus, whilst in Nigeria sows with piglets or immature animals are protected at all times.

 

However, in many African countries, particularly in west and central Africa, law-enforcement is problematic, and in several countries it is virtually non-existent. In Benin, Central African Republic, Gabon, Ghana, Nigeria, Sierra Leone, Liberia, Sudan and Zaire (now Democratic Republic of Congo), for example, poaching is rife even within many nominally protected areas. By comparison, wildlife protection laws in some other countries, such as Zimbabwe, Botswana, Namibia and South Africa, are sufficiently well enough enforced to largely preclude the poaching of these animals within protected areas, though farmers may still be permitted to exercise control of their numbers elsewhere.

 

 

Captive Breeding

 

Both species are extremely rare in captivity outside their countries of origin, though specimens have been exhibited at intervals in various collections in Europe and the U.S.A. since the middle of the 19th century. However, they have been bred only very occasionally, and never in sufficient numbers to be self-sustaining, let alone meet the undoubted demand for exhibition purposes. Longevity was also very poor amongst the early captives, though a female red river hog was maintained at the Frankfurt Zoo from 1959 to 1979, whilst another female of the same species currently held in the Duisburg Zoo is close to breaking this 20 year record (Gewalt, 1988).

 

A small group of red river hogs has been maintained and bred in the Duisburg Zoo since 1979, though these animals are all descended from a single wild-caught pair from Togo. These animals, and another wild-caught pair from Togo recently imported by the Los Angeles Zoo, are thought to be the only specimens of this species held anywhere outside Africa at the present time. Moreover, the prospects for any further exports are diminishing with the imposition of increasingly stringent veterinary regulations concerning the movement of wild pigs throughout the European Community and in North America.

 

 

Additional Remarks

 

These species are maintained in a number of collections in Africa. However, few if any efforts have been made to develop properly structured research or captive breeding programmes within their countries of origin, or to explore their evident commercial potential for pork production. Although there is no evidence that Potamochoerus has ever been truly domesticated, Simoons (1953) has cited references to the semi-domestication or at least taming of P. porcus by various peoples along the northern border of the rainforest zone, as well as early accounts which suggest the species was taken to South America, where it was encountered in feral and tamed states, and to England, where it was alleged to have been used for cross-breeding with domestic pigs, S. scrofa (see earlier text). The (presumed) introduction of P. larvatus to Madagascar and the Comoro Is. might also reflect an earlier, and hitherto largely unsuspected, cultural and economic importance attached to these animals amongst some ethnic groups, which undoubtedly merits further investigation. In the Iyiocha Forest Reserve in southern Nigeria, for example, local tribal groups accord P. porcus the status of "king of the forest" (W. Oduro, pers. comm.). If an animal is killed, its carcass is carried to the traditional chieftain's palace before it is allowed to be butchered, and the meat is shared among the members of the chief's clan, and only a portion is given to the hunter.

 

 

Conservation Measures Proposed:

An Action Plan

 

Given that both species remain widely distributed, are well represented in numerous national parks and reserves, and are maintaining their numbers outside these areas in many parts of their ranges, neither species should be regarded as threatened at the present time. Indeed, in some areas there may even be a need to instigate more rigorous population control measures, as in some parts of Madagascar where (introduced?) bushpigs are reported to pose a serious threat to some native wildlife species. However, whilst species' conservation per se is not an issue at present, these species are threatened in some countries/regions, and efforts should be made to conserve these populations. In addition, very few field studies have been conducted to date, and many basic aspects of their systematics, biology and management requirements are poorly known and merit further investigation.

 

Objectives:

 

1. To obtain a better understanding of the distribution, genetic variation and systematic relationships of these species.

 

2. To obtain distribution and population status data in those areas where this information is lacking at present.

 

3. To encourage further studies of the population biology, ecology and behaviour of these species, and their socio-economic importance to tribal societies.

 

4. To promote the development of management programmes designed to: a) protect remnant populations in countries/regions where either species are threatened, or b) ensure the sustainable harvesting of these animals in areas where they remain abundant as an important resource for local people.

 

 

Priority Projects:

 

 1. Obtain additional cranial, skin and cytogenetic specimen material (and photographs) from particular key locations, with a view to the resolution of outstanding systematic questions.

 

Various questions pertaining to the geographic and genetic variation in these species, and their systematic relationships, remain unresolved at present owing to the absence or shortage of comparative specimens from particular key locations. These locations and questions include: a) Madagascar: further research is needed to check the validity of the subspecies hova and larvatus, and the relationships (and possible origins) between these animals and those of the African mainland; b) east and south Zaire and neighboring countries: obtain comparative specimens in the contact zone between P. porcus and P. larvatus, investigate reports of polymorphism in some of these populations, and reassess the distribution and taxonomic status of these animals in the light of these results; and c) Somalia and north-east Kenya: obtain additional, comparative taxonomic materials to assess the validity of separating 'P. l. somaliensis' from P. l. hassama, and obtain more details about the distribution and status of bushpigs in this region. (Also see Grubb, this vol.).

 

2. Promote field studies on the biology and ecology of P. larvatus and, particularly P. porcus.

 

Very few field studies have been conducted on either of these species and, as far as is known only one such study (in southern Nigeria by Oduro, 1989) has been undertaken on P. porcus. Further, comparative studies are required in order to obtain a proper understanding of their natural history and possible future management needs.

 

3. Collect population distribution and status data in selected areas.

 

At present, available data are inadequate or entirely lacking from a number of regions/countries, including Sudan, northern Ethiopia, south-east Somalia, central Zaire, Togo, Angola and Mozambique. In some of these areas, however, continued political unrest may preclude data collection in the near future, and may be a cause for concern in respect of its implications for increased hunting pressure to provide bush meat to military and displaced persons, and the ineffective protection of designated parks and reserves.

 

4. Investigate occurrence and status of introduced and feral pig populations, encourage their control or eradication, and discourage any future introduction attempts.

 

Updated data are required on the distribution and status of all naturalized pig populations, and the threats these pose to the genetic integrity and health of native wild pigs and other species. In particular, recent reports of P. porcus X S. scrofa hybrids in Gabon and elsewhere should be investigated, and voucher specimens of presumed hybrids (skeletons, skins, blood and cell samples) should be obtained because of their scientific interest. Nonetheless, support should also be given to the control or removal of these animals, just as similar support should be given to the control of any introduced populations of P. larvatus in those locations in Madagascar or the Comoro Islands where they are known or presumed to pose a primary threat to native wildlife species. (Also see Oliver and Brisbin).

 

5. Encourage anthropological (including archaeozoological) research on the socio-economic significance of Potamochoerus and other native wild pigs amongst ethnic groups, and promote studies designed to investigate the possible future use of these animals for domestic husbandry. (Also see Oliver et al. this vol., Chapter 4.9).

 

 

Acknowledgements

 

Grateful thanks are extended to the following correspondents who completed the African Suiform Survey report forms and supplied numerous reprints and other data: F. O. Amubode, W. F. N. Ansell, J. J. Ballestra, R. H. V. Bell, A. Blom, J. C. Bolibar, T. Butynski, A. Camara, P. Chardonnet, B. Child, G. Davies, A. Dupuy, F. Fagotto, J. M. Fay, J. M. Fryxell, M. E. J. Gore, J. M. Grandvergne, A. A. Green, R. Harding, J. Hart, J. C. Hillman, K. Hillman-Smith, N. Innocent, R. C. V. Jeffery, A. Laurent, L. Macky, H. Mertens, J. Maronga-Mbina, N. Monfort, F. E. C. Munyenyembe, J. Newby, M. Nicoll, B. Y. Ofori-Frimpong, A. Peal, F. Rakotondraparany, P. J. J. van Rensburg, C. A. Spinage, M. Storz, G. Teleki, J. P. Thomassey, P. Trenchard, J. Verschuren, S. Vrahimis, P. L. Walker and V. Wilson. Peter Grubb and Kristin Leus provided numerous valuable comments and suggestions on earlier drafts of this manuscript, and Peter Cuypers was largely responsible for the preparation of the distribution map.

 

 

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