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

 

Introduction

 

The Old World pigs and their New World analogues, the peccaries, are important to people. Hippos are also important but, being (spectacularly) more specialized, much less so. Nonetheless, the arrival of human settlers on Madagascar is implicated in the extinction of up to three species of endemic hippopotamuses in the last one and a half millennia (Grubb, this vol., section 4.1). Elsewhere human exploitation of the suiformes dates back for tens of thousands of years. In many parts of the world, wild pigs have long constituted the most commonly eaten large animal. The earliest human remains (c. 40,000 yrs. BP) in the Niah Caves of Sarawak (Borneo) are associated with large numbers of bearded pig (S. barbatus) bones (Cranbrook, 1979), and recent studies of the economy of hunted species in this country have demonstrated that this species is still by far the most important game animal (Caldecott and Nyanoi, 1985).

 

Pigs were also amongst the first species to be domesticated, and it is now certain that this happened in several different places and times with different (local) progenitors. Most of these progenitors were regional variants of the Eurasian wild pig (Sus scrofa) but, as Groves (1981) revealed, the Sulawesi warty pig (S. celebensis) has also been domesticated, and its purebred, hybrid and feral derivatives are still hunted and husbanded in various parts of South-east Asia and the Papuan Realm. In these regions, in particular, the economic importance of wild and domestic pigs led inevitably to their being interwoven with the cultures of the people who depended upon them. Surviving examples of these 'pig cultures' are still to be found amongst some tribal groups in these areas, and it is apparent that pigs are as fundamental to the integrity and economy of these societies as cattle are to other, comparable societies in the drier regions of the Indian Subcontinent and Africa.

 

Being highly adaptable, more prolific and more suited to backyard and small-scale commercial husbandry, pigs are also more abundant and more widely distributed than other hoofstock and they are still amongst the most important of all domestic animals. In Asia alone, the annual consumption of pork had exceeded 20 million tons per year by the mid-1980's, an amount greater than the total consumption of all other domesticated species put together (FAO, 1985).

 

The importance of these animals as a basic food resource was also reflected in their widespread carriage and dispersal during the early transmigrations of settling peoples, who either released them to be hunted whenever required for eating or who maintained them in varying states of domestication.

 

Their dispersal was continued by the later European explorers, sealers, whalers, immigrant settlers and colonialists, and most recently (and irresponsibly) by commercial and recreational hunters and game meat producers; all of whom have transported and (whether accidentally or deliberately) released founder stocks to form naturalized populations. As a result, pigs are now one of the most widely distributed of all species, and the diversity of wild and domestic, feral, hybrid, native and introduced forms, has produced patterns of distribution and interrelationships of almost unparalleled confusion.

 

This complexity of forms is most apparent in South-east Asia, where an inordinate number of taxa have been described in the scientific literature. The challenge of unraveling their relationships was taken up by Colin Groves, whose major reviews of Babyrousa (1980) and, especially, Sus (1981) are the basis for our present understanding of the diversity and regional genetic variation in these genera. In clarifying the affinities and distribution of innumerable naturalized populations, many of which were erroneously recognized as valid taxa, Groves also helped to clear the dross and provide a contextual framework for the interpretation and weighting of other, conservation-related data.

 

The current and equally important review of the subgeneric taxonomy of the Afrotropical suids by Peter Grubb (section 4.1, this vol.) has also provided an indispensable rationale for weighting future research and conservation priorities amongst these animals. In this instance, however, the prioritization process was greatly facilitated by the simultaneous collection of questionnaire data on the present distribution and conservation status of all suiform species from most (sub-Saharan) African countries. In 1988/89, following a format similar to that devised by the Antelope Specialist Group (East, 1988), approximately 600 questionnaires (one questionnaire for each species of wild pig and hippo known or believed likely to occur in each country) were sent to a total of 115 wildlife officials and biologists in 42 countries. A total of 236 (39%) completed questionnaires and/or copies of relevant reports, reprints and maps, were returned by 93 (81%) correspondents, with information from 34 (81%) countries. Analyses of these data, together with information gleaned from a variety of other sources, including Grubb's review, provide the basis for the 'action plan' chapters for each species/genus in this volume. A similar questionnaire survey is now being undertaken for the peccaries in Central and South America and, in 1990, in collaboration with the Deer Specialist Group, a questionnaire survey was initiated for all Eurasian and South-east Asian suids. Unfortunately, however, only a very small number of completed questionnaires have been returned to date, though the few data obtained from these have also been incorporated in the relevant species' chapters.

 

The obvious importance of utilizing both latest taxonomic and recent field status data sets in the formulation of conservation plans is exemplified by Grubb's (section 4.1, this vol.) revelations about the warthogs of Somalia and northern Kenya. These animals are diagnostically distinguished from all other living warthogs (P. africanus) by a variety of dental and cranio-morphometric characters. However, these characters are shared by another recent warthog, the so-called 'Cape' warthog (P. aethiopicus), which was known only from the vicinity of Cape Province, where it is now extinct. Clearly, the recognition of the survival of P. aethiopicus elsewhere, and the apparent isolation of the Somali population, can only profoundly influence our perceptions about the relative importance of these animals, in both scientific and conservation terms.

 

Unfortunately, the import of Grubb's review may be bedeviled by bickering between the traditionalists and the revisionists or, as with Phacochoerus, between neontologists and paleontologists, as to the validity of the two species approach. This also applies to the separation of the bushpig and the red river hog as two, distinct species, Potamochoerus larvatus and P. porcus, respectively. Neither of these divisions is new, since both have been proposed by various earlier authors, though never in the light of so comprehensive a re-examination of available museum specimens as that undertaken by Grubb. Despite this, the recognition of two species in each of these genera has already ruffled a few traditionalist feathers, and it seems likely the debate will rumble on as a futile distraction from the more pressing business of addressing the future research and management requirements of these animals.

 

This is not to suggest that Grubb's review, or those of Groves (1980, 1981) before him, are the last word on the subgeneric taxonomy of these animals. Taxonomy is at best an inexact science, and the present subdivisions are no more than a logical reflection of current understanding of the variation within these genera. By the same token, the relative extent of variation within these or other genera, will not only determine where the lines are drawn, but influence the taxonomic ranking accorded to the various subgroupings. Thus, the distinction between species and subspecies is frequently blurred and, in the absence of any precise knowledge of the degree of genetic variation within the genus as a whole, and an objective overview of that variation, it may become nonsensical to dismiss 'subspecies' as intrinsically less important than 'species'.

 

These sentiments can be illustrated by the recent 'elevation' of two 'new' species of Sus, both endemic to the Philippines. These are the Visayan warty pig (S. cebifrons) from the West-Central Visayas Islands and the East Philippines' warty pig (S. philippensis) from Luzon, Mindanao and associated islands. Following Sanborn (1970), both of these taxa were assigned to S. celebensis until Groves (1981) reassigned them as endemic subspecies of S. barbatus. However, following the acquisition of additional skulls and mandibles in 1990, Groves (1991) suggested these were actually sufficiently distinct to merit full species status; a view which has recently (too recently for the full results to have been incorporated in this volume) been strongly endorsed by de Haan et al. (in press) on account of their lower chromosome number (i.e. 36, as opposed to the 38 typical of other members of the genus) and other cytological characters.

 

The addition of these two species, together with at least one endemic and one endemic subspecies of S. barbatus (i.e. ahoenobarbus from Palawan and associated islands, and barbatus from Tawi-tawi and Sibutu, respectively), makes the Philippines the second most important country in the world (after Indonesia) for its diversity of wild pigs. There may even be still more taxa awaiting description in the archipelago. S. philippensis, for instance, is currently regarded as monotypic, though the few adult individuals which have been examined from the two, principal populations on Luzon and Mindanao, appear morphologically distinct in some respects (e.g. color and shape of the crest and mandibular warts, color of pelage on cheeks, etc.), and these may be new subspecies. Theoretically, the wild pigs of Mindoro should be closely related to those of neighbouring Luzon, though the only specimen examined to date shares the same chromosome number (38) as those from Palawan (de Haan et al., in press), which it also resembles more closely in appearance. Wild pigs have already been extirpated on two (Tablas and Romblon) of the three islands in the isolated Romblon Group, and although they are reported to survive on Sibuyan, no museum specimens exist. This situation also applies to Masbate and a number of other, relatively isolated islands where pigs are reported to survive, as well as to Siquijor where they are now extinct (Cox, 1987; Oliver, 1992; Oliver et al., this vol., Fig. 14).

 

In any event, it is clear that much more work is needed in countries like the Philippines, where we have only very recently begun to obtain an understanding of the genetic diversity and future management requirements of these animals. Moreover, some of this work is undoubtedly urgent. Whatever was on Siquijor has been lost, and whatever the identity of the wild pigs on Sibuyan, Masbate, Bohol and many other smaller, or more isolated, islands, these populations are seriously threatened or likely to become so in the near future. The recently recognized Visayan warty pig is already extinct on two islands (Cebu and Guimaras) and endangered on the only other islands (Negros and Panay) on which it is known to occur. As a result, this species is now rated as one of the most endangered of all wild suids.

 

In a very real sense, this situation also reflects a much wider, and in many ways quite astonishing, upheaval in our understanding and appreciation of the genetic diversity and conservation status of the suiformes which has occurred over the past twenty years or so. The key events include the recognition, in 1969, of an isolated population of pygmy hippo from the Niger Delta, as a separate subspecies (Hexaprotodon liberiensis heslopi). The present status of this animal is not known, though it is rumored to survive (Corbett, 1969; Grubb, this vol. a; Eltringham, this vol.). In 1971, the (somewhat dramatized) 'reappearance' of the pigmy hog (S. salvanius) was announced by tea planters in N. W. Assam, following the burning of grasslands in one of only two small areas still known to support remnant populations of this critically endangered species. The continuing tenuous existence of this species, arguably one of the most important indicator species for the enhanced management of the crucially important tall grasslands, and certainly one of the world's potentially most valuable genetic resources, is still the cas celebre of the Pigs and Peccaries Group, but by no means its only concern. One of the most amazing large mammal discoveries of this century, that of the giant peccary (Catagonus wagneri), has also emerged as one of this Group's major preoccupations (Taber, this vol.). C. wagneri was originally described in the 1930's from late Pleistocene material, and its survival was unknown until some fresh skulls were collected from local hunters during mammal inventory project in the Chaco of Paraguay in 1972 (Wetzel et al., 1975).

 

In the process of unraveling the genus Sus, Groves (1981) also unearthed a 'mystery pig' (S. bucculentus), which is still known from only two skulls collected in Cochin China (southern Vietnam) before the turn of the century, and described a new subspecies of warty pig (S. v. blouchi) from skulls recently collected from the tiny island of Bawean, off N. E. Java. Despite its minuscule range, the Bawean warty is thought to be relatively secure at present, though the animal's external appearance remains virtually unknown. Thanks to the observations of Hoogerwerf (1970) in Udjong Kulon National Park, and various earlier authors, much more information was available on the closely related Javan warty pig (S. v. verrucosus). However, these animals inexplicably disappeared from Udjong Kulon sometime during the 1970's and, in the absence of contemporary reports of their occurrence elsewhere, they were feared extinct in the wild by the end of the decade (J. MacKinnon, pers. comm.). Fortunately, this proved not to be the case, though the implementation of recommendations arising from an island-wide status survey conducted the following year at the Group's behest, have proved problematic (Blouch et al., 1983; Blouch, this vol.).

 

A rather similar situation obtained at about this time with another, equally extraordinary suid, the 'golden' or 'hairy' babirusa (B. b. babyrussa), from Buru and the Sula Islands. However, it was not until the late 1980's, after an interval of about thirty years, that the first confirmed reports of the continuing existence of these animals were finally obtained (Macdonald, this vol.). As in this instance, relevant field data may be very difficult to acquire simply because of the remoteness or inaccessibility of the areas. Buru, for example, being a penal colony until quite recently, was closed to visitors. In countries like Indonesia and the Philippines, the sheer logistics of trying to obtain recent population status data, let alone research specimens, over all or most of a species' known or presumed range, is at best daunting and at worst either effectively impossible or cost prohibitive. Despite a high probability of new taxa awaiting discovery, or becoming extinct before discovery, many islands have never been surveyed or even visited by biologists. Permits for obtaining specimens for captive breeding programs or scientific study, and/or accessing those specimens for examination by specialists, is also becoming increasingly difficult. This is particularly true in the case of wild pigs, where concern (sometimes amounting to paranoia) in the pork industry about the possibility of disease transmission, has resulted in total bans or excessively stringent veterinary regulations being imposed on the movement of wild pig specimens across (in some cases even within) national boundaries.

 

Nonetheless, this work continues as opportunities are presented or created, and a good deal of progress has been made in some areas. Even so, the exercise is often difficult, frequently more frustrating than rewarding, and almost always salutary. Even as initial priorities are identified and (where practical) addressed, old problems are seldom vanquished and new questions, problems and priorities multiply. The recent upheaval in our understanding of these animals is far more a reflection of our ignorance than of our knowledge. A great deal of basic data on the distribution, population status, ecology and behavior of most species of wild suiformes are still required, and these needs must be tackled if we hope to develop the most appropriate management strategies for these species.

 

In the broadest terms, these are the reasons and the justification for this Action Plan - or, more precisely, these 'action plans', since the modest number of extant species (currently 18, excluding the enigmatic Sus bucculentus) within the purview of the Pigs and Peccaries and the Hippo Specialist Groups, has enabled us to produce an 'action plan' for each species, and even a few related topics. The format adopted (which has been unashamedly lifted from the now, sadly defunct, Red Data Books) has also allowed us the luxury of including summaries of much of what is already known about the taxonomy, distribution, ecology and behavior of each species. This was done both as a deliberate attempt to pull these data together, and as a means of identifying some of the starkest gaps in our knowledge; thereby providing perspectives and justifications for most of the emergent recommendations. By the same token, we have endeavored to produce priority recommendations for useful future research and conservation management of each of these species and topics, both as an exercise for people with specialist interests and as the source of reference for the various, highest priority recommendations which are brought together in the regional reviews at the end of each section.

 

William L. R. Oliver

 

 

References

Anon 1993. First Cypriots feasted on hippos. New Scientist 137 (1855): 18.

 

Blouch, R. A. 1993. The Javan warty pig, Sus verrucosus. pp 129-136. In: Pigs, Peccaries and Hippos Status Survey and Action Plan. Ed. by W.L.R. Oliver. IUCN/SSC Pigs and Peccaries Specialist Group and IUCN/SSC Hippos Specialist Group, Gland, Switzerland 202 pp.

 

Blouch, R. A., Banjar Y. L., Heri D. S., and Sumaryoto A. 1983. The Javan Warty Pig: Distribution, Status and Prospects for the Future. (Unpubl.) rep. to World Wildlife Fund, Bogor: 32pp.

 

Caldecott, J. and Nyaoi, A. 1985. Sarawak's wildlife: a resource to be taken seriously. Sarawak Gazette April 1985: 31-32.

 

Corbet, G. B. 1969. The taxonomic status of the pygmy hippopotamus Choeropsis liberiensis, from the Niger Delta. J. Zool. 158: 387-394.

 

Cox, C. R. 1987. The Philippine spotted deer and the Visayan warty pig. Oryx 21 (1): 37-42.

 

Cranbrook, Earl of, 1979. A review of domestic pig remains from archaeological sites in Sarawak. Sarawak Mus. J. XXVII (48): 79-87.

 

East, R. 1988. Antelopes. Global Survey and Regional Action Plans, Part 1: East and Northeast Africa. IUCN, Gland: 96 pp.

 

Eltringham, S. K. 1993.: The pygmy hippopotamus, Hexaprotodon liberiensis. pp. 55-60. In: Pigs, Peccaries and Hippos Status Survey and Action Plan. Ed. by W.L.R. Oliver. IUCN/SSC Pigs and Peccaries Specialist Group and IUCN/SSC Hippos Specialist Group, Gland, Switzerland 202 pp.

 

FAO. 1985. Production yearbook, 1984. Food and Agriculture Organisation, Rome.

 

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 Prehistory, Research School of Pacific Studies, Australian National University, Canberra: 96 pp.

 

Groves, C. P. 1991. Wild pig skulls from Mt. Talinis, S. E. Negros. (Unpubl.) rep. to British Museum (Natural History): 2 pp.

 

Groves, C. P. and Grubb, P. 1993. The Eurasian suids, Sus and Babyrousa - taxonomy and description. pp 107-111. In: Pigs, Peccaries and Hippos Status Survey and Action Plan. Ed. by W.L.R. Oliver. IUCN/SSC Pigs and Peccaries Specialist Group and IUCN/SSC Hippos Specialist Group, Gland, Switzerland 202 pp.

 

Grubb, P. 1993a. The Afrotropical hippopotamuses, Hippopotamus and Hexaprotodon - taxonomy and description. pp 41-43. In: Pigs, Peccaries and Hippos Status Survey and Action Plan. Ed. by W.L.R. Oliver. IUCN/SSC Pigs and Peccaries Specialist Group and IUCN/SSC Hippos Specialist Group, Gland, Switzerland 202 pp.

 

Grubb, P. 1993b. The Afro-tropical suids, Phacochoerus, Hylochoerus and Potamochoerus - taxonomy and description. pp 66-75. In: Pigs, Peccaries and Hippos Status Survey and Action Plan. Ed. by W.L.R. Oliver. IUCN/SSC Pigs and Peccaries Specialist Group and IUCN/SSC Hippos Specialist Group, Gland, Switzerland 202 pp.

 

de Haan, N. A., Bosma, A. A., Macdonald, A. A. and Oliver, W. L. R. 1992. A new species of wild pig in the Philippines with a type of centric fusion translocation new to Sus. Proc. 10th European Colloquium on Cytogentics of Domestic Animals, Utrecht.

 

Hoogerwerf, A. 1970. Udjung Kulon, the Land of the Last Javan Rhinoceros. E. J. Brill, Leiden: pp 331-349.

 

Macdonald, A. A. 1993. The babirusa, Babyrousa babyrussa. pp 161-171. In: Pigs, Peccaries and Hippos Status Survey and Action Plan. Ed. by W.L.R. Oliver. IUCN/SSC Pigs and Peccaries Specialist Group and IUCN/SSC Hippos Specialist Group, Gland, Switzerland 202 pp.

 

Oliver, W. L. R. 1992. The taxonomy, distribution and status of Philippine wild pigs. Silliman J. 36 (1): 55-64.

 

Oliver, W. L. R., Cox, C. R. and Groves, C. P. 1993. The Philippine Warty Pigs, Sus philippensis and S. cebifrons. pp. 145-155. In: Pigs, Peccaries and Hippos Status Survey and Action Plan. Ed. by W.L.R. Oliver. IUCN/SSC Pigs and Peccaries Specialist Group and IUCN/SSC Hippos Specialist Group, Gland, Switzerland 202 pp.

 

Wetzel, R. M., Dubos, R. E., Martin, R. L. and Myers, P. 1975. Catagonus, an 'extinct' peccary alive in Paraguay. Science, 189: 379-381.

 

 

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