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The Suborder Suiformes consists of threeextant families: the Suidae, Dicotylidae andHippopotamidae. These are now thought tocomprise at least 19 living species, of which allbut three species- the collared peccary Tayassutajacu, the red river hog Potamochoerus porcusand the bush pig P. larvatus - are threatened tovarying degrees, either throughout their rangesor because they include one or more threate-ned subspecies. Even the widely distributed andoften locally abundant Eurasian wild pig (Susscrofa), common warthog (Phacochoerus africa-nus) and white-lipped peccary (T. pecari) allhave threatened subspecies, which meritincreased conservation attention. By compari-

son, the status of a few taxa, such as the pigmyhog (S. salvanius), the recently recognisedVisayan warty pig (S. cebifrons) and theNigerian race of pigmy hippo (Hexaprotodonivoriensis heslopi), are critical; whilst others, likethe enigmatic Vietnam warty pig (S. bucculen-tus), may already have followed the three spe-cies of Madagascar dwarf hippos (Hippopotamuslaloumena, H. lemerlei and Hexaprotodon mada-gascariensis) and the Cape race of the “desert”warthog (P. a. aethiopicus) into extinction.Recent revisions of the taxonomy of these ani-mals are summarized in table 1 and the presentknown conservation status of all currentlyrecognized taxa are summarized in table 2.

3IBEX J.M.E. 3:1995

TAXONOMY AND CONSERVATION STATUS OF THE SUIFORMES - ANOVERVIEW

Oliver W.L.R.IUCN/SSC Pigs and Peccaries Specialist Group. Park End, 28A Eaton Road, Norwich, Norfolk NR4 6PZ, U.K.

Keywords: Suidae, Dicotylidae, Hippopotamidae, Endangered species, Wild pigs.

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Order: Artiodactyla (comprising three suborders: Suiformes, Tylopoda, and Ruminantia)Suborder: Suiformes (comprising two superfamilies and three families, as follows):

Superfamily: Anthracotheroidea

Family: Hippopotamidae(no separate subfamilies, but two genera and two living and three recently extinct species, as follows):

Genus: Hippopotamus Species: H. amphibius (5 ? ssp) - Common HippoH. laloumena - Madagascan Hippo (extinct)H. lemerlei - Madagascan Dwarf Hippo (extinct)

Genus: Hexaprotodon Species: H. liberiensis (2ssp) - Pygmy Hippo(=Choeropsis) H. madagascariensis - Madagascan Pygmy Hippo (extinct)

SuperFamily: Suoidea (comprising two families, seven genera and ≤ seventeen species, as follows):

Family: Dicotylidae (no separate subfamilies, but two genera and three species, as follows):

Genus: Tayassu Species: T. tajacu (14 ? ssp) - Collared PeccaryT. pecari (c. 5 ssp) - White-lipped Peccary

Genus: Catagonus Species: C. wagneri (0 ssp) - Giant or Chacoan Peccary

Table 1. Taxonomy of living and recent suiformes.

Total: superfamilies 2, families 3, subfamilies 3, genera 9, species (provisionally 22 (of which 3, possibly 4, are extinct), sub-species (provisionally) > 65 (of which at least 1, but possibly 2 or more, are extinct).

Oliver W.L.R. Status of species, genetics and conservation - Communication

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Taxon CurrentConservation Status

Family Hippopotamidae

Hippopotamus amphibius 2*Common Hippopotamus(subspecific taxonomy requires review)

Hippopotamus laloumena ExtinctMadagascan Hippopotamus

Hippopotamus lemerlei ExtinctMadagascan Dwarf Hippopotamus

Hexaprotodon liberiensisPygmy Hippopotamus

H. l. liberiensis 4H. l. heslopi Indeterm. (prob.6)

Hexaprotodon madagascariensis ExtinctMadagascan Pygmy Hippopotamus

Family Dicotylidae

Tayassu tajacu 1-2*Collared Peccary(subspecific taxonomy requires review)

Tayassu pecariWhite-lipped Peccary

T. p. ringens 3T. p. spiradens 3 - 4T. p. equatorius Indeterm.T. p. pecari 1 - 2T. p. albirostris 1

Family: Suidae (comprising three subfamilies, five genera and ≤ fourteen species, as follows):

Subfamily: Suinae - the “true” pigs

Genus: Sus Species: S. scrofa (c. 17 ssp) - Eurasian Wild PigS. salvanius (0 ssp) - Pygmy HogS. bucculentus - Vietnam Warty Pig (extinct ?)S. verrucosus (2 ssp) - Javan Warty PigS. barbatus (3 ssp) - Bearded PigS. cebifrons (0 ? ssp) - Visayan Warty PigS. philippensis (? ssp) - Philippine Warty PigS. celebensis (0 ? ssp) - Sulawesi Warty Pig

Genus: Potamochoerus Species: P. larvatus (>3 + ? ssp) - BushpigP. porcus (0 ssp) - Red River Hog

Genus: Hylochoerus Species: H. meinertzhageni (4 ssp)- Forest Hog

Subfamily: Phacochoerinae - the warthogs

Genus: Phacochoerus Species: P. aethiopicus (2 ssp, 1 extinct) - Desert WarthogP. africanus (c. 4 ssp) - Common Warthog

Subfamily: Babirousinae - babirusa

Genus: Babyrousa Species: B. babyrussa (4 ? ssp, 1 possibly extinct) - Babirusa

Table 2. Summary of present conservation status of Suiformes.

Taxon CurrentConservation status

Catagonus wagneri 5Chacoan PeccaryFamily Suidae

Phacochoerus aethiopicusDesert Warthog

P. a. aethiopicus ExtinctP. a. delamerei** 4

Phacochoerus africanusCommon Warthog(subspecific taxonomy provisional)

P. a. africanus 1P. a. aeliani Indeterm. (prob. 5)P. a. massaicus 1P. a. sundevallii 1 - 2

Hylochoerus meinertzhageniForest Hog

H. m. meinertzhageni 3 - 4H. m. rimator 3 - 5H. m. ivoriensis 5H. m. ssp. (S. Ethiopia) Indeterm.

Potamochoerus larvatusBushpig (subspecific taxonomy provisional)

P. l. hassama 2P. l. (?) somaliensis 2P. l. koiropotamus** 1P. l. larvatus (introduced ?) 1 - 2P. l. hova (introduced ?) 1 - 2

Status of species, genetics and conservation - Communication Oliver W.L.R.

5IBEX J.M.E. 3:1995

Key:

* = Future taxonomic reviews likely to result in one ormore subspecies being included in a more threatenedcategory.

** = Additional taxonomic material from selected areaslikely to indicate taxon comprises/includes two ormore subspecies.

Status category definitions:

1 = ‘Widespread and abundant’;

2 = ‘Known or believed relatively secure’ (i.e. wide-spread at low density, but abundant in some areas orlimited distribution but abundant and not thought tobe threatened, and/or well represented in protectedareas);

3 = ‘Potentially at risk or Rare’ (i.e not thought to beimmediately threatened, but has restricted distributionor is widespread but nowhere abundant, but occurs insome protected areas);

4 = ‘Known to be at risk or Vulnerable’ (i.e. hasrestricted distribution an/or limited ecological tole-

rance, known to be threatened by habitat destruc-tion/disturbance, hunting pressure or other factors,over the majority of its range, and/or is inadequatelyrepresented in reserves);

5 = ‘Seriously threatened or Endangered’ (i.e. highlyrestricted and/or fragmented distribution; all knownpopulation declining and status likely to become criti-cal in the near future);

6 = ‘Critically endangered’ (i.e. only one or a very few,small populations, which are unlikely to surviveunless urgent action is taken to redress causative fac-tors);

Extinct = ‘Extinct’.Indeterm. = Indeterminate: ‘Taxa considered to be

Threatened’ (i.e. status categories 3 to 6 above, butavailable data insufficient to determine appropriatecategorisation.

REFERENCES:All data modified afterOLIVER W.L.R. (ed.) 1993: Pigs, Peccaries and Hippos:

Status Survey and Conservation Action Plan.IUCN, Cambridge and Gland, 202 pp.

(Family Suidae, continued)

Potamochoerus porcus 1Red River Hog

Sus scrofaEurasian Wild Pig

S. s. scrofa 1S. s. meridionalis 2 - 3S. s. algira 2S. s. attila 1S. s. lybicus 1 - 2S. s. nigripes 1S. s. davidi 1S. s. cristatus 1S. s. affinis 1S. s. ssp. (Bopeta, Sri Lanka) Indeterm.S. s. sibiricus 1S. s. ussuricus 1S. s. leucomystax 2S. s. riukiuanus 4 - 5S. s. taivanus 2 - 3S. s. moupinensis 1S. s. vittatus 1

Sus salvanius 6Pygmy Hog

Sus bucculentus Extinct ?Vietnam Warty Pig

Sus verrucosusJavan Warty Pig

S. v. verrucosus 4S. v. blouchi 4

Sus barbatusBearded Pig

S. b. barbatus 2S. b. oi 3S. b. ahoenobarbus 3

Sus cebifrons 5 - 6**Visayan Warty Pig

Sus philippensis 3**Philippine Warty Pig

Sus celebensis 1 - 2**Sulawesi Warty Pig

Babyrousa babyrussaBabirusa

B. b. babyrussa 5B. b. celebensis 4B. b. togeanensis 5B. b. bolabatuensis Indeterm. (Extinct?)

6 IBEX J.M.E. 3:1995

CONSERVATION GENETICS OF THE GENUS Sus

Randi E.Istituto Nazionale per la Fauna Selvatica, via Cà Fornacetta 9, 40064 Ozzano dell’Emilia, Bologna, ltaly.

1. IntroductionThe Suidae comprise a number of species ofgreat biological interest and economical value.Suids interact with human activities, becausethey are domesticated, reared, crossed, translo-cated, hunted, eated, and in certain cases,venerated or persecuted. They have a place inmany traditional cultures and in the everydaylife of millions of humans. Such notwithstan-ding, the biology of many species is still poorlyknown, and their management and conserva-tion cannot take advantage of the necessarybasic information.Conservation genetics, the application ofpopulation (and in particular small popula-tions) genetic models and biochemical andmolecular technologies to conservation bio-logy, is aimed to preserve biodiversity at thedifferent levels it is organized (genes, popula-tions, species, evolutionary lineages), and canbe applied to address a wide range of still openquestions about the biology of the Suidae.First of all we need to develop a molecular phy-logeny of the Suidae, which must integratenon-molecular knowledges, and constitute thebackground within which to describe the timescales and patterns of their evolution. This fra-mework is necessary to delineate species (aswell as subspecies and populations) boundariesand relationships. Just to mention a few exam-ples: the wild pig fauna of entire regions (e.g.,the Philippines) is poorly known, andspecies/subspecies boundaries have not beenclearly delineated so far (Oliver et al., 1993).The patterns of speciation and partitions of

genetic diversity among populations of theAfrican suids await description (Grubb, 1993),as well as it is not known the extent of geneticdivergence among the 3 (or more ?) subspeciesof the babirusa (Macdonald, 1993). Captivebreeding is frequently mentioned as a necessaryoption to secure the preservation of someendangered suids (first of all, the highly endan-gered Pygmy hog, Sus salvanius; Oliver & DebRoy, 1993). Captive propagation with the per-spective of reintroduction in the wild, needsthe preservation of the largest possible fractionof the species’ gene pool. Quantification of exi-sting gene diversity within small populations,breeding plans aimed to retain the maximumpossible gene diversity and to avoid inbreedingand inbreeding depression are efforts requiredto reconstitute viable populations in the wild.Suids represent a resource for human popula-tions. Two species have been domesticated(Sus scrofa and S. celebensis; Groves, 1981), andother are easily bred in captivity (NationalResearch Council, 1983). Most populations ofthe Eurasian wild pig are intensively exploitedand managed, but the genetic, demographicand ecological impacts of intense hunting pres-sure and animal translocations are still poorlyknown. In selected cases it will be important todetermine the origins of feral pig populations,and evaluate eventual genetic peculiarities,because, besides the obvious problems theypose as pests in exotic habitats, they couldhave some value for conservation and as possi-ble source of human incomes (Brisbin, 1990),once appropriately managed and controlled.

Keywords: Sus scrofa, Suidae, Molecular biology, Populations, Hybridization, Mitochondrial DNA.

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Abstract: Recent technical advances in biochemical and molecular biology make it possible to study the popula-tion genetics and phylogenetics using biological samples collected undestructively during field researches, or collec-ted from museum specimens. Applications of conservation genetics of the genus Sus range from the definition of amolecular phylogeny of the Suiformes to the description of boundaries between species and subspecies and theidentification of the origins of wild, feral, domestic or cross-breeded populations. In this paper I review informationon the genetic structure of west European wild pig populations, and present recent findings on nucleotide variabi-lity of mitochondrial DNA genes. The amplification of DNA through the polymerase chain reaction and followingnucleotide sequencing of selected mitochondrial DNA genes, are effective methods which can be applied to descri-be patterns of within- and among- species genetic variability in the Suidae.

Status of species, genetics and conservation - Communication Randi E.

7IBEX J.M.E. 3:1995

2. Population genetics of Sus scrofa.Multilocus Protein (enzymes and non-enzyma-tic proteins) Electrophoresis (MPE) is an inex-pensive and simple technique, widely used tostudy population genetics and phylogenetics ofmany animal species. Proteins migrate in anelectric field because some aminoacids arecharged. If a point mutation (or a deletion, aninsertion) changes the aminoacid compositionof a protein, it will change (in about 30% ofthe cases) its net electric charge, and its rate ofelectrophoretic migration will be different anddistinguishable from the original molecularform. Appropriate staining recipes will, then,reveal allelic variation at single structural loci.Therefore, it is possible to identificate andcount genotypes and alleles, compute within-population estimates of gene diversity,between-population genetic distances, andwork up a number of population geneticmodels like Hardy-Weinberg equilibrium,linkage disequilibrium, gene flow, geographicalpopulation structuring, and so forth.Limitations of protein electrophoresis derivefrom the low mutation rates at structural loci,and from possible selective value of some alle-lic variants. lt is not expected any aminoacidsubstitution in populations which were recen-tly separated, and the recovery of heterozygo-sity is slow after a recent bottleneck (Lande &Barrowclough, 1987). In some cases proteinsmay be not sufficiently variable to allow discri-minating among recently isolated populations.Some populations of the Eurasian wild pighave been recently studied, using karyotypemapping (Bosma et al., 1984), blood groups(Kurosawa et al., 1979) and MPE (Tanaka etal., 1983; Randi et al., 1989). Results concor-dantly showed the existence of an east-west cli-nal variation in allele frequencies, which pro-duced dendrograms separating the eastern fromthe western phenotypically recognized subspe-cies (Kurosawa et al., 1979). Studies of restric-tion fragment length polymorphism of themitochondrial DNA (Watanabe et al., 1986;Lan & Shi, 1993) supported these conclusions.The observed genetic gap is roughly correspon-dent to the distributions of the two prevalentkaryotypes, 2n = 36 and 2n = 38 (Bosma et al.,op.cit.). An insufficient sampling of wild pigpopulations along an east-west transect fromcentral Asia to central Europe prevents us tounderstand the origins of these concordant pat-terns of genetic variation: the apparent clinalvariation, and the apparent main genetic gapcould be the results of recent dispersal from dif-

ferent and anciently isolated centers of origins,and/or of isolation by distance acting on popu-lations with restricted (at least across someareas) gene flow. Multivariate morphometricsof skull measures indicated a more or less linearnorth-east to south-west dimensional cline,perhaps mainly due to environmental effectsand adaptations to food availability on growthrates and adult body size (Randi et al., 1989).Blood groups and MPE showed comparativelygreater genetic distances between Asian andEuropean-American domestic pig breeds(Tanaka et al., op.cit.), supporting the idea ofpolyphyletic domestication (Oliver et al.,1993). Some of the modern pig breeds origina-ted from Asian stocks or possibly from crossesbetween Asian and European pig strains(Ollivier & Sellier, 1982). These informationscan be used to investigate the population gene-tic structure and the putative origins of localwild pig populations, particularly in caseswhere translocations and/or introductionscould have originated populations of uncertainorigins or unknown genetic make up. We haveused MPE to describe genetic variability insome west European wild pig populations (S.s.scrofa), and to assess the genetic structure ofthe formerly described ltalian subspecies (S.s.meridionalis, the Sardinian wild pig, and S.s.majori, the Maremma wild pig; Randi et al.,1989). Clustering multilocus pairwise geneticdistances indicated small divergence amongS.s. majori and other west European wild pigpopulations (Fig.1). These results prompted usto reject the validity of the subspecies S.s.majori, which must be considered, at best, anecotype phenotypically adapted to aMediterranean type habitat. Within relativelyshort geographic distances, and in contrastwith the presence of significant body size diffe-rences, allozyme variability is often not structu-red enough to evidence significant populationgaps. We have recently studied genetic variabi-lity in some Bulgarian wild pig populationssampled in localities where were described(Genov et al., 1991) the existence of differentmorphological phenotypes: a larger one livingin the northern plains, and a smaller oneoccurring in the southern mountains. MPEshowed results concordant with morphometrics(Fig. l), but allele divergence among northernand southern populations was small (Randi etal., 1992). Such small distances are not neces-sary attributable to genetic isolation and absen-ce of gene flow, but most probably to geneticdisequilibrium and drift, with consequent ran-

Randi E. Status of species, genetics and conservation - Communication

8 IBEX J.M.E. 3:1995

dom fluctuations in allele frequencies, due tothe genetically small effective size of recentlyestablished populations. We observed greatergenetic distance values separating wild andferal Sardinian pigs from other west Europeanwild pig populations, thus validating the sub-species S.s. meridionalis (Fig. l). The locus6PGD is a possible marker of hybridizationbetween wild and domestic pigs. This locus waspolymorphic in almost all the studied pigbreeds (Franceschi & Ollivier, 1981), while itwas monomorphic in all the studied westEuropean wild pig populations (Randi et al.,1989). It was monomorphic in Sardinian ferals,as well as in the Smoky Mountains pigs(USA), which originated, at least in part, fromintroduced European wild pigs (Smith et al.,1980).We hypothesized that 6PGD is polymorphic inAsian wild pigs and that this polymorphismwas introduced in modern pig breeds throughhybridization (Randi et al., 1989). Thishypothesis has been supported by Kurosawaand Tanaka’s paper (1991) showing extensive6PGD polymorphism in S.s. leucomystax andS.s. riukiuanus (with some monomorphic popu-lations, possibly due to recent bottlenecks andisolation), and S.s. taivanus. 6PGD and otherallozyme polymorphisms (PGM, PGI) are

linked to artificially selected meat productioncharacters (e.g., in the linkage group PGI-HAL-6PGD; Rasmusen, 1983), and can showallele frequency divergence among wild anddomestic pigs. These enzyme loci could be usedto locate the geographic origins of certain wildpig stocks, and to detect hybridization andintrogression of domestic genes at the popula-tion level.

3. Mitochondrial DNA nucleotide sequen-cingUsing Restriction Fragment LengthPolymorphisms (RFLP) we can estimate ratesof nucleotide substitutions through restrictionendonuclease digestions of purified targetDNA, followed by detection of restriction frag-ments by hybridization with a cloned labelledprobe. The main advantage of this method (aswell as of most DNA methods) is given by thepossibility to choose the appropriate targetDNA sequence to be studied. DNA is not ahomogeneous strip of nucleotides, but is astructured macromolecule with single copystructural genes and repetitive DNA, withgenes evolving at a low pace, or hot spots ofmutation and recombination, i.e., sequenceswith high rates of molecular evolution. Usingcloned hypervariable sequences as probes, it is

Figure 1 - UPGMA dendrogram obtained by clustering allozyme Nei’s genetic distances among some west European wild,feral and domestic pig populations.


9IBEX J.M.E. 3:1995

possible to identificate the single individual(DNA fingerprinting), while using slowly evol-ving sequences as probes (e.g., RNA genes), itis possible to study phylogenetic relationshipsamong very divergent taxa. The main draw-back of RFLP method is the need of good qua-lity and abundant target DNA, which cannotso easily be obtained during field work or unde-structively collected from endangered species.Recently direct nucleotide sequencing hasbecome feasible at the population level, thanksto the discovery of the possibility to amplificateDNA using the polymerase chain reaction(PCR). The target DNA sequence is amplifiedin vitro using two oligonucleotide primers(synthesized in vitro). One primer is comple-mentary to the sequence flanking one end ofthe target DNA, while the other primer iscomplementary to the other flanking sequence.The presence of free nucleotides and of a ther-mostable DNA polymerase, results in theextension of both primers, which copy the tar-get DNA sequence. PCR is performed in anautomated machine, the thermal cycler, whichcontrols a 3-temperature cycle: DNA denatura-tion, primer hybridization, DNA extension.This cycle can be repeated 20-50 times, dou-bling the quantity of target DNA each cycle,and producing, at the end, a million times theamount of target sequence present initially.This pure DNA can be directly sequenced.Nucleotide sequences produce enormousamounts of genetic information: the singlenucleotide is the character (with four possiblestates), and sequences are exactly comparablethrough different laboratories. Sequences arecumulative information, and DNA databasesare exponentially growing. One of the most informative application ofboth RFLP and PCR in population and conser-vation genetics is the study of mitochondrialDNA (mtDNA). MtDNA is present in themitochondria, it is maternally inherited(through the oocytes), it is haploid and doesnot recombine. MtDNA shows (at least inmammals) an average rate of molecular evolu-tion 5-10 times faster than average single copynuclear DNA. This makes mtDNA the mole-cule of choice to study genetic divergenceamong conspecific populations, and to deter-mine maternal phylogenies. We are running aproject to sequence selected mtDNA genes inthe Suidae, with the aim: 1) to identificatereliable slow-evolving regions, which canretain unambiguous phylogenetic signals andwhich can allow reconstructing the evolutio-

nary patterns of taxa within the Suiformes; 2)to identificate fast-evolving hypervariableregions, which can be used to describe patternsof genetic variability at the interface betweenconspecific and intraspecific populations. Wehave designed oligonucleotide PCR primers toamplificate and to sequence the entire mito-chondrial cytochrome b (CYB), a protein-coding gene, with intermediate rate of sequen-ce evolution in mammals, and the mitochon-drial control region (D-LOOP), a non-codingregion involved in the replication of themtDNA, and usually evolving at high rate inmammals. We have sequenced about 600 bp ofthe CYB and 500 bp of the D-LOOP in severalspecies of the Suidae and in samples of diffe-rent populations of west European wild pig.The main results (Fig. 2) indicate that: 1) CYB showed low sequence variation in S.scrofa, and divergence among species was dueto point mutations only. Preliminar calibrationof the CYB molecular clock agrees with presu-med paleontological information, and othermolecular findings, on times of speciationwithin the Suiformes, and suggests that CYB isa suitable gene to obtain reliable phylogeneticsignals at the ordinal level. 2) D-LOOPshowed extensive sequence reorganizationamong different species, due to both pointmutations and duplications of repeated motifs,as well as to insertion or delection of singlebases. Therefore, D-LOOP seems to be anunreliable source of phylogenetic informationin the Suidae. West European wild pig samplesshowed a surprisingly low level of sequencevariation at the D-LOOP. Most populationsshared the same (CYB + D-LOOP) mtDNAhaplotype, including domestic pigs, and someSardinian wild and feral pigs. Sequence diver-gence within west European S. scrofa (wild anddomestic) is less than 1%, and we have notfound any mtDNA fixed sequence differencebetween wild and western domestic pigs.About 60% of the studied Sardinian wild pigspecimens showed a different mtDNA haploty-pe (0.3% nucleotide divergence from the com-mon haplotype), which supports the indica-tions of genetic peculiarity of the Sardinian pigpopulation, and further confirms his subspecificstatus.

4. ConclusionsThe analysis of patterns of nucleotide sequencedivergence, both among- and within-species, isa preliminar step, necessary to identificatemtDNA genes which could be used as a relia-


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ble source of information for intraordinal phy-logenetics or population genetics studies in theSuiformes. The mtDNA CYB is a protein-coding gene, and as expected it is structurallyconserved across species, as consequence ofevolutionary constraints. In the SuiformesCYB seems to evolve at approximately 2%nucleotide substitutions per million year,accordingly with the average rate of mtDNAmolecular evolution in mammals. Our findingssuggest that CYB can be reliably used to descri-be times of evolution and pattern of phyloge-netic divergence in the Suiformes. CYB will beparticularly useful to individuate phylogeneticgaps due to pre- or post-Pleistocene isolation ofevolutionary lineages, and could be successfullyapplied to disentangle species relationships inareas, like the Philippines, of particular biogeo-graphic complexity. The D-LOOP appears toevolve at a faster rate in the Suiformes. lt ismainly a non-transcribed region, with lowerconstraints to both genome size and structuralreorganization. Correct alignment of reorgani-zed D-LOOP regions could be difficult inanciently separated taxa, and consequently theD-LOOP could be reliable to track phylogenyat lower ranks only.Some portions of the D-LOOP should behypervariable within species, and therefore

should be informative for the description ofphylogeographic relationships among conspeci-fic populations or clusters of closely related spe-cies. The 500 bp of the D-LOOP we havesequenced in west European wild pig samples,so far, showed a surprisingly low rate of nucleo-tide variability. Genetic variability amongpopulations was concordantly low at nuclearallozyme loci, mtDNA CYB, as well as atmtDNA D-LOOP, supporting the idea of arecent colonization of western Europe by S.scrofa populations which survived a relativelyintense bottleneck. These findings furtherlystress the existence of high phenotypical plasti-city in S. scrofa, which favours great adaptabi-lity to very different habitat and food condi-tions, and suggest caution in using morphome-try as a tool for intraspecific taxonomy. Withinsuch a framework, it is noteworthy the presen-ce of a peculiar mtDNA haplotype in about60% of the studied Sardinian wild pigs. Pigs (aswell as most of the present large vertebrates)were most probably introduced by man inSardinia (Groves, 1989). The observed geneticheterogeneity of the Sardinian pig populationcould have been originated from multipleintroductions of pigs of different geographicorigins. The peculiar mtDNA Sardinianhaplotype has not been detected in any other

Figure 2 - Neighbor-joining tree showing phylogenetic relationships among Suiformes, obtained using nucleotide sequencesof the mitochondrial DNA cytochrome b gene.


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west European wild pig population, so far. ltcould represent the original mtDNA haplotypeintroduced in Sardinia, probably through semi-domesticated pigs from the Middle East. Theother mtDNA haplotype detected in theSardinian pig population is the common westEuropean pig mtDNA haplotype, which couldhave been introduced in Sardinia later. Thishypothesis points out to two problems. First ofall we wish to track the geographic origin anddistribution of the Sardinian mtDNA haploty-pe, with a particular attention to the status ofthe wild pigs of Corsica and Andalusia, whichare currently described as S.s. meridionalis(Groves, 1981). The second one has more rele-vant implications for conservation. TheSardinian pig population is a genetically mixedone. lt is probable that the actual Sardinian pigpopulation is a patchwork of ancient feral pigs(probably originated in the Middle East),which have been introgressed with recentwestern domestic pigs. If so, we need to analysethis patchwork, mapping haplotypes occurenceand frequencies, and enforce the preservationof ancient Sardinian pigs (the true S.s. meridio-nalis) in the areas which will be eventuallydiscovered to be unpolluted.Concordantly with karyotypes, protein markersand mtDNA RFLP, we expect that mtDNAnucleotide sequences will show greater diver-gence among Asian and European wild pigpopulations, as well as domestic pig breeds(work in progress). The individuation of fixedmtDNA sequence differences between diffe-rent wild pig populations, or between wild anddomestic pig breeds will greatly aid the analysisof the genetic origins of managed wild pigpopulations, as well as the individuation ofwild x domestic crosses.MtDNA is maternally inherited and can cha-racterize maternal ancestries only. PCR can beused to perform a very sensitive fingerprintinganalysis through the amplification of microsa-tellites. Microsatellites consist of 10-50 copiesof 1 to 6 bp repeats, which are randomly inter-spersed in all eucariotic DNAs. There is anenormous number of microsatellites and, asmany microsatellites have 4 or more alleles ofdifferent length (generated probably by asym-metrical crossing-over), they can describe anenormous amount of variability. These repeatsare flanked by DNA with unique sequences,which can be used for locus-specific priming forPCR amplification. Microsatellites can be usedas fingerprinting, to detect hybrids if parentalsare fixed for different alleles, and for popula-

tion genetics, if population divergence hasbeen very recent. We can take advantage ofthe great number (probably more than 400)microsatellite loci which have been describedin the domestic pig (Rohrer et. al.,1994), andthat work perfectly on wild pigs’ DNA (unpu-blished results). One of the most important advantages of PCRis that it makes possible to amplify very smallquantities of DNA (in theory starting from asingle DNA molecule), also if recovered invery bad conditions. DNA suitable for PCRcan be obtained from almost any kind of smallsamples (one drop of blood, one hair root, afew square millimeters of skin biopsy), also if invery bad state of preservation (old museumskins, old bones, archaeological remains). Freshand old samples can be preserved at room tem-perature in absolute ethanol, without necessityto frozen them. It is therefore possible to col-lect samples for PCR amplification and DNAsequencing as by-product of many field workprojects, as well as from museum specimens.

5. Acknowledgements I wish to thank A. L. Bolelli, P. De Marta andV. Lucchini for assistance during laboratoryanalyses at Istituto Nazionale per la FaunaSelvatica, and C. H. Diong (Singapore), P.Genov (Sofia), G. Massei (Firenze), G. Tosi(Milano), A. Zilio (Milano) for collaborativeprojects and sample collection.

REFERENCESBOSMA A. A., DE HAAN N. A. & MACDONALD A. A.,

(1984) - Karyotype variability in the Wild boar(Sus scrofa). Symposium International sur leSanglier. F. Spitz & D. Pépin (eds), Toulouse, LesColloques de l’INRA, n° 22: 53-56.

BRISBIN I. L. JR., (1990) - A consideration of feralswine (Sus scrofa) as a component of conservationconcerns and research priorities for the Suidae.Bongo, 18: 283-293.

FRANCESCHI P. F. & OLLIVIER L., (1981) - Fréquencesde quelques gènes importants dans les populationporcines. Z. Tierzuchtg. Zuchtgsbiol., 98: 176-186.

GENOV P., NIKOLOV H., MASSEI G. & GERASIMOV S.,(1991) - Craniometrical analysis of BulgarianWild boar (Sus scrofa) populations. J. Zool.,Lond., 225: 309-322.

GROVES C. P., (1981) - Ancestors for the pigs:Taxonomy and phylogeny of the genus Sus.Technical Bull. No 3. Department of Prehistory,Research School of Pacific Studies, AustralianNational University, Canberra.

GROVES C.P., (1989) - Feral mammals of theMediterranean islands: documents of early dome-stication. In: The walking larder: patterns of domesti-


12 IBEX J.M.E. 3:1995

cation, pastoralism and predation. J.Clutton-Brock(ed.), Umvin Hayman, London: 46-58.

GRUBB P., (1993) - The Afrotropical suids (Phaco-choe-rus, Hylochoerus and Potamochoerus). Taxonomyand description. In: Pigs, Peccaries and Hippos. W.L. R. Oliver (ed), IUCN, Gland, Switzerland: 66-75.

KUROSAWA Y., OISHI T., TANAKA K. & SUZUKI S.,(1979) - Immunogenetic studies on wild pigs inJapan. Anim. Blood Grps. Biochem. Genet., 10:227-233.

KUROSAWA Y. & TANAKA K., (1991) - PGD variantsin several wild pig populations of east Asia.Animal Genetics, 22: 357-360.

LAN H. & SHI L., (1993) - The origin and genetic dif-ferentiation of native breeds of pigs in southwe-stern China: an approach from mitochondrialDNA polymorphism. Biochem. Genet., 31: 51-60.

LANDE R. & BARROWCLOUGH G. F., (1987) - Effectivepopulation size, genetic variation, and their use inpopulation management. In: Viable populations forconservation. M. E. Soulè (ed.), CambridgeUniversity Press, Cambridge: 87-123.

MACDONALD A. A., (1993) - The babirusa (Babyrousababyrussa). In: Pigs, Peccaries and Hippos. W. L. R.Oliver (ed.), IUCN, Gland, Switzerland: 161-171.

NATIONAL RESEARCH COUNCIL, (1983) - Little-knownAsian animals with a promising economic future.Washington.

OLIVER W. L. R., COX C. R. & GROVES C. P., (1993) -The Philippine warty pigs. (Sus philippensis and S.cebifrons). In: Pigs, Peccaries and Hippos. W. L. R.Oliver (ed.), IUCN, Gland, Switzerland: 145-155.

OLIVER W. L. R. & DEB ROY S., (1993) - The pygmyhog (Sus salvanius). In: Pigs, Peccaries and Hippos.W. L. R. Oliver (ed.), IUCN, Gland, Switzerland:121-129.

OLIVER W. L. R., GROVES C. P., COX C. R. & BLOUCHR. A., (1993) - Origins and domestication and thepig culture. In: Pigs, Peccaries and Hippos. W. L. R.Oliver (ed.), IUCN, Gland, Switzerland: 171-179.

OLLIVIER L. & SELLIER P., (1982) - Pig genetics: areview. Ann. Genet. Sel. Anim., 14: 481-544.

RANDI E., APOLLONIO M. & TOSO S., (1989) - Thesystematics of some Italian populations of Wildboar (Sus scrofa): a craniometric and electropho-retic analysis. Z. Säugetierk., 54: 40-56.

RANDI E., MASSEI G. & GENOV P., (1992) - Allozymevariability in Bulgarian Wild boar populations.Acta Theriologica, 37: 271-278.

RASMUSEN B. A., (1983) - Isozymes in swine breeding.In: Isozymes, Vol. 11. Alan R. Liss, Inc., NewYork: 249-268.

ROHRER, G. A., ALEXANDER L. J., KEELE J. W., SMITHT. P. & BEATTIE C. W., (1994) - A microsatellitelinkage map of the porcine genome. Genetics, 136:231-245.

SMITH M. W., SMITH M. H. & BRISBIN I. L. JR., (1980)- Genetic variability and domestication in swine.J. Mamm., 61: 39-45.

TANAKA K., OISHI T., KUROSAWA Y. & SUZUKI S.,(1983) - Genetic relationships among several pig

populations in East Asia analysed by blood groupsand serum protein polymorphisms. Anim. BloodGrps. Biochem. Genet., 14: 191-200.

WATANABE T., HAYASHI Y., KIMURA J., YASUDA Y.,SAITOU N., TOMITA T. & OGASAWARA N.,(1986) - Pig mitochondrial DNA: polymorphism,restriction map, orientation and sequence data.Biochem. Genet., 24: 385-396.

1. IntroductionRelatively few data are available on the rela-tionships between morphometrical parametersand biochemical-genetical attributes of Wildboar (Sus scrofa). Mostly the growing patternsof different parameters were analysed and fewresults were presented on the correlations andallometry of these characteristics. Better mana-gement of Wild boar populations requires moredata about the morphological and especiallyabout the genetical characteristics of this spe-cies. For this it is important to map the geneti-cal structure of different free-living stocks andto designate the most valuable ones.The purpose of this study was to evaluate thebiochemical variability and differentiation ofWild boar populations in Hungary. Theresearch program covered the following areas:• Evaluation of biochemical methods for the

analysis of enzyme-polymorphism in Wildboar.

• Search for polymorphic enzymes in Wildboar populations to detect the most poly-morphic enzymes and investigation of fre-quent allele variations. The year-to-yearvariability of enzyme-polymorphism andmorphological parameters were also sur-veyed.

• The relationships between morphologicalparameters (body weight-body length, bodyweight-height at shoulder, body weight-cir-cumference of chest) and comparison of sex-and age-dependent variations were estimatedby statistical methods.

2. Material and MethodsBody weight, body length, height at shoulderand circumference of chest data of shot animals(n=188) from both sexes were collected in the1991/92 and 1992/93 hunting seasons. Heart,liver and kidney tissue samples were taken fromthe animals during evisceration and the sam-ples were stored deep frozen until the labora-tory processing.The following enzymes were investigated:Malic-enzyme (ME, E.C. 1.1.1.40), Isocitrat-dehydrogenase (IDH, E.C. 1.1.1.42), Acid-phosphatase (ACP, E.C. 3.1.3.2), Catalase(CAT, E.C. 1.11.1.6), Hexokinase (HK, E.C.2.7.1.1.), Glucose-dehydrogenase (GDH, E.C.1.1.1.47). Electrophoretic and staining pro-cedures were completed according to routinemethods described in the literature (Hartl &Höger, 1986 modified by Ernhaft, 1991).Polymorphism and heterozygosity were deter-mined by horizontal starch-gel electrophoresisand enzyme-specific procedures for fiveisoenzymes.

3. Results and discussionThe routine electrophoretic and stainingmethods used in Red deer studies proved to beappropriate for Wild boar. Similarly to Reddeer the homogenization could be omittedwhich quickened the laboratory processes.Polymorphic alleles were found in the ME-1,IDH-2, LDH-2, ACP-1, ACP-2 and HK loci.ME-1 and IDH-2 were the most variableisoenzyme loci studied. No polymorphic differ-

13IBEX J.M.E. 3:1995

DATA ON THE BIOCHEMICAL-GENETICAL POLYMORPHISM OF WILDBOAR IN HUNGARY

Ernhaft J., Csányi S.Wildlife Biology Station, University of Agricultural Sciences, H-2103 Gödöllõ, Hungary.

Keywords: Wild boar, Sus scrofa, Suidae, Polymorphism, Enzymes, Isoenzymes, Morphometry, Europe.

IBEX J.M.E. 3: 13-14

Ernhaft J., Csányi S. Status of species, genetics and conservation - Poster

ences were found between the samples of1991/92 and 1992/93 seasons.The most important correlations are presentedin table 1. No morphometrical differences wereshown between the samples of 1991/92 and1992/93 season.On the basis of the isoenzymes the biochemical

REFERENCESHARTL G. & HÖGER H., (1986) - Biochemical varia-

tion in purebred and crossbred strains of domesticrabbits (Oryctolagus cuniculus L.). Genet. Res.(Comb.), 48: 27-34.

ERNHAFT J., (1991) - Biochemical and morphometricaldifferentiation between Red deer (Cervus elaphusL.) and Fallow deer (Dama dama L.) in Hungary.Trans. XXth Congress of IUGB, 21-25 August,Gödöllõ , Hungary, Part 2: 803.

14 IBEX J.M.E. 3:1995

Parameters Slope of regression (b) Correlation coefficient (r)

1991 1992 1991 1992

Body weight - body length 0.53 0.49 0.79 0.74Body weight - height at shoulder 0.20 0.23 0.41 0.48Body weight - circumference at chest 0.46 0.52 0.84 0.80

Table 1. Regression slopes and correlation coefficients for body size parameters.

and genetical characteristics of Wild boar anddomestic pig were also compared. Lowerbiochemical-genetical variability of domesticpigs was found and the allelic variation of ME-1, IDH, GDH, ACP-1 in Wild boar was higherthan in domestic pigs.

1. IntroductionIn Piedmont, following dramatic environmen-tal modifications, the Wild boar diffusedrapidly since the early sixties also as a conse-quence of absence of predators, decrease of cul-tivation, and transformation of cattle, sheepand goats breeding techniques, particularly inmountain zones. Some role is also due to intro-duction for hunting purposes of crosses withdomestic pig and of wild boars from EasternEurope. Therefore we have now a well adaptedanimal which sometimes presents biometricaltraits similar to those of domestic pig. However,biometrical traits, being multifactorial, are onlyin part expression of gene activity, being hea-vily influenced by environment. Instead bio-chemical polymorphisms are useful markers tostudy the genetic structure of a population,because they are controlled by genes on whichthe environmental influence is almost null.

2. Material and methods The samples (liver, heart, kidney and diaph-ragm) come from 197 wild boars hunted during1990-1992 in different mountain areas ofPiedmont and from 26 domestic pigs from aslaughterhouse as a control. By means of hori-zontal starch gel electrophoresis the loci GPI,PGD, LDHA, LDHB, ADA1, ADA2, ME1,ME2, MDH1, MDH2, IDH1, IDH2, SODA,SODB, HK, ACP were analyzed. The resultshave been compared with those relative to Susscrofa scrofa from Austria (Hartl & Csaikl,1987), from peninsular Italy (Apollonio et al.,1985) and with those of Sus scrofa meridionalisfrom Sardinia (Apollonio et al., op. cit.).

3. Results GPI system has been found polymorphic as inAustrian Wild boar (GPI*A .24, GPI*B .76);in domestic pig the frequency of GPI*B variesfrom .30 to .97. PGD system showed a single allele (PGD*A)as it has been observed in other Wild boarpopulations. In domestic pig the system is poly-morphic and the frequency of PGD*A variesfrom 0.30 to 0.83.ADA1 and ME1 were monomorphic in oursample in contrast with Austrian populations.The remaining systems were monomorphic.

4. Discussion and conclusions All the wild boars of the Western Europe so farexamined are similar, although coming fromdifferent zones. The constant trait is the mono-morphism of PGD locus wich is polymorphicin domestic pig and in some populations ofAsian Wild boar (Kurosawa & Tanaka, 1991).This seems to indicate that the alleles PGD*Band PGD*C arose from mutation when Susscrofa scrofa was already differentiated. In thiscase, if in a population of Wild boar one ofthese two alleles should be present, the occu-rence of crosses with domestic pig or of indivi-duals of Asian origin could be hypothesized,although conclusions from a single locus cannot be absolute. However, karyotype analysis(Macchi et al., in press) evidenced that chro-mosome polymorphism is absent in individualsfrom the same mountain zones, where the pre-sent material has been collected. It seems rea-sonable to accept that in the Piedmont moun-tains is present the typical Western Europe

15IBEX J.M.E. 3:1995

GENETIC CHARACTERIZATION OF SOME POPULATIONS OF WILDBOAR (Sus scrofa scrofa) IN PIEDMONT (ITALY)

Durio P. *, Macchi E. **, Rasero R. ** Dip. Produzioni Animali, Epidemiologia ed Ecologia, via Nizza 52, 10126 Torino, Italia. ** C.R.E.A. - Centro Ricerchein Ecologia Applicata, via Catti 12, 10146 Torino, Italy.

Keywords: Wild boar, Sus scrofa scrofa, Suidae, Population genetics, Biochemical genetics, Polymorphism, Europe,GPI, PGD.

IBEX J.M.E. 3:15-16

Abstract: Electrophoretic data were obtained from 200 wild boars hunted in Piedmont mountains and were compa-red with those from other feral populations.

Wild boar (Sus scrofa scrofa), on which intro-duction of crosses or wild boars of other subspe-cies did not have notable influence. Anyway, itseems to be useful to control all the Wild boarcenters existing in Piedmont both withkaryotype and biochemical polymorphism exa-mination because, if differences from wildpopulation should be found, it would be possi-ble to detect at least one of the sources of ille-gal introduction.

ReferencesAPOLLONIO M., RANDI E. & TOSO S., (1985) -

Morphological and biochemical analysis of someitalian populations of Wild boar. IVth InternationalTheriological Congress, Edmonton, Canada, 1985.

HARTL G.B. & CSAIKL F., (1987) - Genetic variabilityand differentiation in wild boars (Sus scrofa ferusL.): comparison of isolated populations. J. Mamm.68: 119-125.

KUROSAWA Y. & TANAKA K., (1991) - PGD variantsin several wild pig populations of East Asia.Animal Genetics, 22: 357-360.

MACCHI E., TARANTOLA M., PERRONE A., PARADISOM.C. & PONZIO G., this volume - Cytogeneticvariability in the Wild boar (Sus scrofa scrofa) inPiedmont (Italy): preliminary data.

Durio P., Macchi E., Rasero R. Status of species, genetics and conservation - Poster

16 IBEX J.M.E. 3:1995

1. IntroductionThe cytogenetic studies point out a chromoso-mal polymorphism in the Wild boar (Bosma etal., 1984; Nombela et al., 1990). The diploidchromosome number of the West Europeansubspecies is 2n=36, the Eastern Europeanpopulations have 2n=38, while in the subspe-cies of Central and Far East Asia it ranges from36 to 38. The domestic pig has always 2n=38and its karyotype is identical to that of wildboar with 2n=38.The aim of this research is to characterize,under the cytogenetic point of view, the popu-lations of Sus scrofa living in Piedmont.

2. Material and methodsWe analyzed 14 blood samples, drawn from thefemoral vein of wild boars trapped and anesthe-tized in mountainous and plain areas, and 4blood samples from pigs.Two methodologies were used: the first is thestandard one on whole blood (Moorhead et al.,1960), the second is derived from the standardmodified by us and using separated blood.Here we describe the latter because it gave bet-ter results.- Separate the blood using 1 ml of Emagel for

1 ml of blood, leave it for 30 minutes.- Centrifuge the supernatant for 10 minutes, at

900 revs per minute (RPM), with the samequantity of RPMI-1640.

- Remove and discard the supernatant with apipette.

- Add 6 ml of RPMI-1640 Dutch modification,2 ml of fetal bovine serum, 0.2 ml of phy-tohaemaglutinin, 0.1 ml of antibiotics and0.1 ml of heparin (100 U/ml).

- Put the cultures in thermostat for 72 hours(37°C).

- 2 hours before the cell collection add 0.3 mlof Vinblastina sulphate (0.5 µg/ml); wait 1hour then add 0.5 ml of Actinomycin D (100µg/ml).

- Centrifuge at 900 RPM for 10 minutes. - Remove and discard the supernatant with a

pipette.- Add a hypotonic solution (KCL 0.075 M)

pre-warmed for 15 minutes at 37°C.- Centrifuge at 900 RPM for 10 minutes,

remove and discard the supernatant with apipette.

- Add 5 ml of Acetic acid (5%).- Leave it for 5 minutes at room temperature.- Centrifuge at 900 RPM for 5 minutes.- Remove and discard the supernatant with a

pipette.- Add 10 ml of fixative (1 part of glacial acetic

acid + 3 parts of methanol).- After 30 minutes centrifuge at 900 RPM for

10 minutes, remove and discard the superna-tant with a pipette, add the fixative, centri-fuge again at 900 RPM for 10 minutes, remo-

17IBEX J.M.E. 3:1995

CYTOGENETIC VARIABILITY IN THE WILD BOAR (Sus scrofa scrofa) INPIEDMONT (ITALY): PRELIMINARY DATA

Macchi E. *, Tarantola M. *, Perrone A. *, Paradiso M.C. **, Ponzio G. **** C.R.E.A - Centro Ricerche in Ecologia Applicata - Via Belfiore 61 bis, 10126 Torino, Italy.** Servizio Universitario di Genetica Medica - USSL VIII, via Santena 19, Torino, Italy.*** Dipartimento di Genetica, Biologia e Chimica Medica - Facoltà di Medicina - Università di Torino, via Santena 19,Torino, Italy.

Keywords: Wild boar, Sus scrofa, Suidae, Europe, Population genetics, Karyotype.

IBEX J.M.E. 3:17-18

Abstract: The authors have carried out a cytogenetic study on the Wild boar (Sus scrofa L.) in Piedmont in order toassess the origin of the population. The existence of some polymorphism in the diploid number indicates thatPiedmont’s animals may have different origins. The modified technique we used proved satisfying.

ve and discard the supernatant with a pipetteand prepare the slides.

The classic banding technique of Caspersson etal. (1970) and Zech (1973) for the Q bandswas used.

3. ResultsKaryological analysis enabled us to show thechromosomal polymorphism represented by 3variants 2n=38, 2n=37, 2n=36.The karyotype 2n=38 consisted of 5 submeta-centric chromosomal pairs (1-5), 2 subacrocen-tic pairs (6-7), 5 metacentric pairs (8-12), 6acrocentric pairs (13-18) and 2 gonosomes(submetacentric X-chromosome and a smallmetacentric Y).The karyotype 2n=37 presented aRobertsonian translocation between a chromo-some 15 and a chromosome 18, that gives asubmetacentric chromosome, while the twohomologous corresponding are free, accordingto Popescu et al. (1980).Finally the karyotype 2n=36 is homozygotic forthe Robertsonian translocation 15-18.Among the 14 samples analysed, the 6 collec-ted in mountainous areas (4 males and 2 fema-les) had 2n=36. Among the 7 coming from flatareas, 3 subjects (2 females and 1 male) had2n=38, 2 males 2n=36 and 2 females 2n=37.All the 4 domestic pigs (3 females and 1 male)had 2n=38.

4. ConclusionsThe diploid chromosomal number in thesubjects coming from mountainous areas is2n=36, in agreement with the results ofMauget et. al.(1984) and Popescu et al.(1980).The chromosomal polymorphism found in thesubjects from the plain area lets us suppose thatthere were in plain arca animals of unknownorigin, i.e. released for restocking.From the obtained results we may concludethat the technique we developed is efficient,thus a wider sample will allow us to obtainmore precise information on the populations ofthis species.Our hypothesis that mountainous areas, wherenatural selection is stronger, prevent the repro-ductive success of reared animals (some ofwhich are illegally released) could be tested inthe future.We seek for a greater collaboration from thepublic administrations in the collection ofother samples and in the control of all Wildboar farming in order to prevent a proliferationof animals with unclear chromosomal set.

The release of such animals in the wild couldcause great negative effects on agriculture andmodify the genetical structure of the truly wildpopulation of Sus scrofa.

REFERENCESBOSMA A.A., DE HAAN N. A. & MACDONALD A.,

(1984) - Karyotype variability in the Wild boar(Sus scrofa). Symposium International sur leSanglier. F. Spitz & D. Pépin (eds), Les Colloquesde l’I.N.R.A., n° 22: 53-56.

CASPERSSON T., ZECH L. & JOHANSSON C., (1970) -Differential binding of alkylating fluorochromes inhuman chromosomes. Exp. Cell. Res., 60: 315-319.

MAUGET R., CAMPAN R., SPITZ F., DARDAILLON M.,JANEAU G. & PÉPIN D., (1984) - Synthèse desconnaissances actuelles sur la biologie du Sanglier,perspectives de recherche. SymposiumInternational sur le Sanglier. F. Spitz & D. Pépin(eds), Les Colloques de l’I.N.R.A., n° 22: 15-56.

MOORHEAD P.S., NOWELL P.C., MELLMAN W.H.,BATTIPS D.M. & HUNGERFORD D.A., (1960) -Chromosome preparations of leukocytes culturedfrom Human peripheral blood. Experim. CellResearch, 20: 613-616.

NOMBELA J.A., MURCIA R.C., ABAIGAR T. & VERICADJ.R., (1990) - Cytogenetic analysis (GTG, CBGand NOR bands) of a Wild boar population (Susscrofa scrofa) with polymorphism in the south-eastof Spain. Genet. Sel. Evol., 22: 1-9.

POPESCU C.P., QUÉRE J.P. & FRANCESCHI P., (1980) -Observations chromosomiques chez le Sanglierfrancais (Sus scrofa scrofa). Ann. Gen. Sel. anim.,12: 395-400.

ZECH L., (1973) - Fluorescence banding techniques. In:Chromosomes identification, T. Caspersson & L.Zech (eds), Nobel symposia 23, Medicine andnatural science. N.Y., Academic press.

Status of species, genetics and conservation - Poster

18 IBEX J.M.E. 3:1995

Macchi E., Tarantola M. et al.

THE PIG MITOCHONDRIAL GENOME

Hecht W., Dzapo V.Department of Veterinary Genetics, Institute of Animal Breeding and Genetics, Justus Liebig University, Hofmannstr. 10,D-35492 Giessen, Germany.

1. IntroductionMitochondria of animal cells contain an auto-nomous genetic system. The genome itself is adouble stranded circular DNA moleculeapproximately 16-17 kilobase pairs long inmost higher vertebrates. Its gene order andcontent is highly conserved among mammals.This organelle genome possesses the geneticinformation for two ribosomal RNAs, 22tRNAs and 13 proteincoding genes. All genesare tightly packed with no or only a few basesbetween them as spacers. The only major non-coding region is the so called D-loop, locatedbetween the genes for the tRNAs forPhenylalanine and Proline. In this region con-trol elements for the transcription and transla-tion processes are located. The origin for L-strand replication however is placed within acluster of tRNA genes between the genes forCox1 and ND2. Mitochondrial DNA is widelyused to infer phylogenetic relationships andvariability patterns among populations. Herewe report on preliminary data from a sequen-cing project, designed to determine the com-plete nucleotide sequence of the pig mitochon-drial genome.

2. Material and MethodsMitochondrial DNA (mtDNA) from a singleanimal has been purified and cloned into pUCvectors. Recombinant clones were propagatedin E. coli and sequenced by the dideoxymethod, either using the sequenase kit (USB)or by an automated procedure on an ABIsequencer. Gap filling was accomplished bysynthesizing oligonucleotides as sequencingprimers. Restriction enzyme analysis was usedto refine a previously published restriction map(Watanabe et al., 1985) and to search forRFLPs among 42 animals of different origin,

including animals of Asian maternal origin,European wild boars and different domesticbreeds. All methods were performed accordingto standard procedures or suppliers instruc-tions.

3. Results and DiscussionThe pig mitochondrial genome is approxima-tely 16,750 base pairs long. Up to now we havesequenced 15,722 bases and report on analysisof up to 13,674. Complete nucleotide sequen-ces have been determined for the followinggenes: ATPase subunits 6 and 8,Cytochromeoxidase subunits 1, 2 and 3,NADH dehydrogenase subunits 1, 4 and 6, 12sribosomal subunit, tRNAs for Arg, Asp, FMet,Glu, Gly, Ileu, Leu, Lys, Ser, Trp, Tyr, Val.Homology comparisons to sequences fromother vertebrates (Anderson et al., 1981; Bibbet al., 1981; Anderson et al., 1982; Desjardins& Morais, 1990) demonstrate that the pigmitochondrial genome exhibits the same geneorder and content as other mammals. Resultsof homology comparisons are depicted in table1. Homology is highest between pig and cow,whatever subgroup of sequences is compared.Differences in similarity between pig andmouse and pig and man are marginal.Sequence homology between pig and chickenis lowest, which is not astonishing, as thechicken has to be regarded as an outgroupamong this species. As expected, the higherconservation of aminoacid sequences as com-pared to nucleotide sequences is due to theredundancy of the genetic code especially forthe third codon position. This is demonstratedfor the pair pig/cow in table 1.Despite the high similarity between cow andpig sequences, the pig genome is roughly 400base pairs longer. This is due to the presence of

19IBEX J.M.E. 3:1995

Keywords: Pig, Wild boar, Sus scrofa, Suidae, Mammals, Mitochondrial DNA.

Abstract: Restriction analyses, cloning and partial sequencing of pig mitochondrial DNA were performed.Restriction data confirm the previously described differentiation of Asian and European mtDNA types and demon-strate the presence of Asian type mtDNA in one European breed. Samples from wild boars show the same restric-tion patterns as European domestic pigs.

IBEX J.M.E. 3:19-20

additional sequences in the D-loop of the pig.A part of the pig D-loop region consists of atandem repeat of the sequence CGTGCGTA-CA. This is a purine/pyrimidine alteration,characteristic for Z-DNA. Our data confirmthe presence of this tandem repeat, which is sofar unique among mammals except the rabbit(Mignotte et al., 1990), whose D-loop issequenced. Putative promotor and/or signalsequences have also been assigned to certainpositions in the D-loop. Additionally, we iden-tified a sequence as possible origin of L-strandreplication by homology analysis. The sequen-ce reads CTCCCGCCGCAGGAAAAAA-AAGGCGGGAG. Position 22 to 29 is aninverted repeat of positions 1 to 8. This can beregarded as characteristic for loop formingstructures, while the inverted repeat forms astem, the sequences spacing them from the sin-gle stranded loop. This structure is thought tobe created, when in the process of H-strandreplication the L-strand becomes singlestran-ded at that position. The loop could then actas signal sequence or substrate for a factor ini-tialising L-strand replication.Our restriction enzyme analyses confirm thepreviously reported differentiation of Asianand European mtDNA types (Watanabe et al.,op. cit.) and demonstrate the presence of Asiantype mtDNA in the Hampshire breed. Furtherthe Belgian Landrace displays a polymorphicHincII site. Sequence comparisons among pigsyield 98.2% homology for the cytochrome bgene between the gene sequence published byIrwin et al. (1991) and our data.

4. AcknowledgementsWe thank Ms H. Schomber for excellent tech-nical assistance. This work was supported by agrant from the DFG.

REFERENCESANDERSON S., BANKIER A.T., BARRELL B.G., DE BRUIJN

M.H.L., COULSON A.R., DROUIN J., EPERON L.C.,NIERLICH D.P., ROE B.A., SANGER F., SCHREIERP.H., SMITH A.J.H., STADEN R. & YOUNG I.G.,(1981) - Sequence and organisation of the humanmitochondrial genome. Nature, 290: 457-465.

ANDERSON S., DE BRUIJN M.H.L., COULSON A.R.,EPERON L.C., SANGER F. & YOUNG I.G., (1982) -Complete sequence of bovine mitochondrial DNA- Conserved features of the mammalian mitochon-drial genome. J. Mol. Biol., 156: 683-717.

BIBB M.J., VAN ETTEN R.A., WRIGHT C.T., WALBERGM.W. & CLAYTON D.A., (1981) - Sequence andgene organization of mouse mitochondrial DNA.Cell, 26: 167-180.

DESJARDINS P. & MORAIS R., (1990) - Sequence andgene organization of the chicken mitochondrialgenome. A novel gene order in higher vertebrates.J. Mol. Biol., 212: 599-634.

IRWIN D.M., KOCHER T.D. & WILSON A.C., (1991) -Evolution of the cytochrome b gene of mammals.J .Mol. Evol., 32(2): 128-144.

MIGNOTTE F., GUERIDE M., CHAMPAGNE A.M. &MOUNOLOU J.C., (1990) - Direct repeats in thenon-coding region of rabbit mitochondrial DNA -Involvement in the generation of intra- individualand inter-individual heterogeneity. Eur. J.Biochem., 194(2): 561-571.

WATANABE T., HAYASHI Y., OGASAWARA N. &TOMOITA T., (1985) - Polymorphism of mito-chondrial DNA in pigs based on restriction endo-nuclease patterns. Biochemical Genetics, 23(1/2):105-113.

Hecht W., Dzapo V. Status of species, genetics and conservation - Poster

20 IBEX J.M.E. 3:1995

Pig Cow Mouse Man Chicken

Total Sequence 78.2 73.5 70.9 63.2(13,674 bases)

Proteingenes 80.0 74.8 73.3 69.0(7 genes)

12sRNA gene 80.9 74.2 75.1 66.1

tRNA genes 86.3 81.8 81.7 72.1(12 genes)

Aminoacidsequences 89.5(8 genes)

Codonposition 1 88.5

2 95.8

3 54.2

Table 1: Homology comparisons between pig and other vertebrates.

1. The situation in the regionThe eight suid species and numerous subspeciesnative to South and Southeast Asia constitutethe highest diversity within this family foundin any region of the world. Groves (1981) hasgone a long way toward clarifying the taxo-nomy of these animals, but recent reviews ofthe Philippine pigs (Oliver et al., 1993; Groves& Grubb, 1993) have shown that further taxo-nomic revisions are required as new materialbecomes available. With few exceptions, wehave little detailed knowledge of the naturalhistory of suids in the region, although duringthe past decade field surveys have begun toprovide a better idea of the distribution, status,and habitat preferences of many of the taxa.An International Union for Conservation ofNature and Natural Resources (IUCN) actionplan drawing together the existing knowledgeand proposing conservation strategies (Oliver,1993) is the source of much of the data summa-rized in the present paper. Several of the nativesuid species and subspecies have been classifiedas threatened and require conservation actionto save them from extinction. Because thesethreatened taxa occur in a wide variety of habi-tats and interact with many varied human cul-tures, each faces a unique set of problems.Nevertheless, uncontrolled hunting and habi-tat destruction have been identified as the twopredominant threats. Consequently, in mostsituations creation and management of protec-ted areas combined with development andenforcement of sound hunting laws are solu-tions requiring high priority.Understanding what needs to be done is amajor step toward successful conservationaction. Unfortunately a series of factors oftenmake it difficult for the necessary policies to beimplemented.

Perhaps the most important constraint is thelimited funds which governments of the regionare able or willing to commit to the conserva-tion of protected areas and wildlife in general.The World Bank calculates that direct govern-ment funding for protected area managementin all of Asia is about US $ 30-35 millionannually, roughly one third of the amountIUCN estimates is needed as a bare minimumto support only the routine expenditures(Braatz, 1992). As a percentage of total annualnational budgets, funding for protected areamanagement is especially low in those coun-tries most important for suid conservation. Forthe Philippines the figure is only 0.01%, thelowest in Asia. India and Indonesia providetheir parks with only slightly higher propor-tions of 0.03% and 0.06% respectively. By wayof comparison, Bhutan with a figure of 0.29%ranks highest in Asia in this respect. In many societies negative attitudes towardpigs make it difficult to garner public andgovernment support for their conservation. Allpigs are despised and avoided by devoutMuslims who refuse to eat or even touch anypart of the animals, and Islam is the predomi-nant religion in Indonesia and Malaysia.Additionally, farmers who suffer losses fromcrop raiding pigs are understandably reluctantto heed laws aimed at conserving species theyregard as pests.A somewhat paradoxical complication for suidconservation is that wild pigs are often animportant protein source for indigenous socie-ties which obtain much of their subsistencefrom hunting and gathering in the forests. Thismay endear the pigs to conservationists whosee them as ideal sustainable “non-timberforest products”. But where governments aremore interested in converting the forest dwel-

21IBEX J.M.E. 3:1995

CONSERVATION AND RESEARCH PRIORITIES FOR THREATENED SUIDSOF SOUTH AND SOUTHEAST ASIA

Blouch R. A.Coordinator for South and Southeast Asia IUCN/SSC Pigs and Peccaries Specialist Group. Department of Wildlife andNational Parks, Km 10 Jalan Cheras, 56100, Kuala Lumpur, Malaysia.

Keywords: Suidae, Sus spp., Babyrousa babyrussa, Asia, Conservation.

IBEX J.M.E. 3:21-25

lers to plantation workers or agriculturalistsliving in permanent villages, conserving wildpig populations for hunting is not a priority. Tosurvive by hunting wild animals is viewed asbackward, and, in the short term, there aremore profitable ways to exploit forests than tomaintain them as support systems for a relati-vely few people who want to continue with adisappearing way of life. As conservationists we must understand theseattitudes and monetary constraints and searchfor ways to overcome them, or to at least mini-mize them in each situation. Among otherapproaches, this may involve public education,training of local wildlife conservation profes-sionals, encouraging conservation of biodiver-sity rather than species’ conservation, promo-ting controlled hunting of wild pigs, or captivebreeding. In addition, basic ecological researchon virtually all threatened suid taxa is still nee-ded. Much of the funding for these activitieswill need to be sought from international aidagencies and non-governmental organizations.

2. Threatened native suids (Tab.1)

genetic contamination through contact withfree ranging domesticates. On Iriomote Islandthere are plans to build a road through a natio-nal park, increasing access by poachers whohad already reduced pig numbers by half sincethe park was created.Due to widespread governmental and localattitudes ranging from ambivalence to hostilitytoward the pigs, the most effective way to con-serve them is by supporting existing or propo-sed programs directed toward the conservationof biodiversity. In addition, preliminary orrepeat status surveys are needed on all sixislands in the Ryukyu chain.

Sus salvanius: Critically Endangered

The Pygmy hog (S. salvanius) is now known toexist in only two isolated populations in thetall grasslands of northwestern Assam, India,and is considered to be one of the most endan-gered of all mammals. Its extremely reducedbody size (males weigh only about 8.5 kg)makes it potentially a highly valuable geneticresource. The continuing decline in Pygmy hog

Blouch R. A. Status of species, genetics and conservation - Communication

22 IBEX J.M.E. 3:1995

Taxon/Status ThreatsSus scrofa riukiuanus (V or E) Hunting (pest/food), Disease, Genetic contaminationSus salvanius (Critically E) Habitat loss, Hunting (food), Political unrestSus v. verrucosus (V) Poisoning, Hunting (pest/food)Sus v. blouchi (V) Hunting (pest)Sus barbatus oi (R) Habitat lossSus b. ahoenobarbus (V) Hab. loss, Hunting (pest/food)Sus philippensis (R) Hab. loss, Hunting (pest/food)Sus cebifrons (E) Hab. loss, Hunting (pest/food)Babyrousa babyrussa (V or E) Hunting (food), Habitat loss

E = endangered; V= vulnerable; R= rare

Table 1: Threatened Southeast and South Asian Suids

Sus scrofa riukiuanus: Vulnerable or Endan-gered according to populationS. s. riukiuanus, the smallest and the onlythreatened subspecies of Sus scrofa, is endemicto the Ryukyu Islands south of the main islandsof Japan. Numbers are declining rapidly, largelyas a result of overhunting, and it is thought tobe endangered on at least four of the six islandsof the Ryukyu chain. Pigs are killed both forconsumption and as agricultural pests, andcommercial traders export carcasses to gourmetmarkets in Osaka. The subspecies is also threa-tened by a severe skin disease which has spreadthroughout the population on one island, and

numbers is attributable to loss of its grasslandhabitat to human settlements, agriculturalencroachment, commercial forestry, and floodcontrol schemes. Annual burning of the tallgrasses concentrates the hogs in the smallremaining unburned areas where they are parti-cularly vulnerable to hunters. The survival ofthe species is crucially dependent on the inte-grity of the Manas Wildlife Sanctuary, which isunder threat from high numbers of illegalimmigrants and an armed rebellion of local tri-bals.The strategy for the conservation of the Pygmyhog places paramount priority on promoting

whatever actions are necessary to restore andmaintain the Manas Wildlife Sanctuary and itsbuffer reserves. Field surveys of areas known orsuspected to harbor pygmy hogs need to beundertaken, and detailed studies are needed onthe behavior and ecology of the species to esta-blish management criteria. A properly structu-red captive breeding program should be establi-shed to provide animals for eventual reintro-ductions to protected sites within its recentknown habitat.

Sus verrucosus verrucosus: Vulnerable

The Javan warty pig (S. v. verrucosus) is confi-ned to the island of Java where it is sympatricwith the Indonesian subspecies of the Eurasianwild pig (S. scrofa vittatus). It occurs at altitu-des below 800 m and prefers disturbed habitatsand teak plantations to closed canopy forests.Although not uncommon in some areas, theremaining Javan warties are in isolated popula-tions and are subjected to uncontrolled hun-ting and, in some cases, poisoning. They arekilled both by sport hunters and by farmersprotecting their crops. There is evidence thathybridization with S. s. vittatus occasionallyoccurs in the wild, but at current levels it isprobably not a threat to the genetic integrity ofthe species. Javan warty pigs are poorly represented in exi-sting protected areas, and proposals to createthree new nature reserves and expand two exi-sting reserves of importance to the taxon needto be implemented soon. Poisoning must bestopped, and surveys of the extent of markethunting should be undertaken with the objec-tive of formulating means to regulate or elimi-nate the practice. Sport hunters from the citiesprovide a source of income to rural peopleacting as guides, and Javan warty populationsoutside of protected areas should be managedto allow this activity on a sustainable basis.Captive animals need to be administered undera properly structured plan for the long termgenetic and demographic benefit of the species.

Sus verrucosus blouchi: Vulnerable

S. v. blouchi, a smaller race of the Javan wartypig than the nominate form, is confined to the200 sq km Bawean Island in the Java Seawhere it is the only suid present. The Muslimpopulation of the island does not hunt the pigsfor food, but crop raiders are snared and killed.Much of the habitat of this subspecies consists

of forests and teak plantations located within a4,500 ha wildlife reserve which was created pri-marily for the benefit of the endemic deer Axiskuhli. But because of its very restricted rangethe subspecies cannot be considered secure,and a survey to determine its current status andconservation requirements is urgently needed.

Sus barbatus oi: Rare

This race of the bearded pig is confined toPeninsular Malaysia and the island of Sumatrain Indonesia where it is sympatric with S. scro-fa vittatus. It is known to travel over greatdistances in large herds of sometimes hundredsof individuals, and was widespread throughoutthe formerly unbroken rain forests characteri-stic of its range. Today forest fragmentation hasdisrupted these movements and reduced thesubspecies’ numbers. For example S. b. oi hasbeen extirpated from the southern end ofSumatra, the portion of the island where defo-restation has been heaviest during the past 40years. These habitat changes have almost cer-tainly favored scrofa over barbatus. Huntingdoes occur, but apparently not intensivelyenough to be a major threat. In PeninsularMalaysia, sport hunters are said to prefer thetaste of Eurasian wild pigs to that of beardedpigs (Pan Khang Aun, pers. comm.). The sub-species is found in several protected areas inSumatra and in the Taman Negara NationalPark in Peninsular Malaysia. Current knowled-ge of S. b. oi is too limited to enable formula-tion of practical management recommenda-tions, and priority must be given to field sur-veys and other basic research to determinewhere they still occur, where and why theymigrate, and whether they can survive in log-ged over forests. A properly structured captivebreeding program should also be initiated.

Sus barbatus ahoenobarbus: Vulnerable

The Palawan bearded pig (S. b. ahoenobarbus)is endemic to Palawan and the smaller islandsof Balabac and the Calamians in thePhilippines. This pig is not known to under-take the long migrations for which the othertwo races of bearded pig are famous. Because ithas a restricted range undergoing rapid defore-station, is known to be found in only one smallprotected area, and is subjected to heavy hun-ting pressure, the Palawan bearded pig is themost threatened race of its species. The fieldstatus survey recently initiated in the Calamian

Status of species, genetics and conservation - Communication Blouch R. A.

23IBEX J.M.E. 3:1995

Islands needs to be continued throughout therange of the race with a view to the develop-ment of recommendations for the enhancedfuture protection of selected populations.Management of the pigs in non-protected areasshould be designed to enable their continuedharvest on a sustainable basis.

Sus philippensis: Rare

The Philippine warty pig (Sus philippensis) isfairly widespread in most of the few remainingforests on the larger islands of the easternPhilippines. However deforestation coupledwith uncontrolled hunting have already extir-pated it over a large proportion of its formerrange and continue to threaten the species. Asa first step to its conservation, field surveys andresearch into habitat requirements, populationdynamics, and response to hunting pressureand commercial logging are needed. Local con-servation education projects should be imple-mented to make the public aware of the impor-tance of their native wildlife and forests ingeneral, and their wild pigs in particular.

Sus cebifrons: Endangered

This most threatened of the Philippine suids,the Visayan warty pig (S. cebifrons), has beeneliminated from four of the six islands where itwas known to exist. On Negros and Panay itsurvives in a few small, isolated populationswhich are still hunted intensively. Forestdestruction continues to reduce and fragmenttheir remaining habitat. The overwhelmingpriority for the conservation of this species isthe early declaration and effective future pro-tection of the proposed Panay MountainsNational Park; other smaller areas with rem-nant populations should also be given protec-ted status. This can be done in the broadercontext of protecting the whole range of criti-cally threatened species endemic to theVisayan faunal region. Data on the present sta-tus of the species are required from severalislands. A properly structured captive breedingprogram needs to be developed, and publiceducation projects should be initiated to raiselocal people’s awareness of their natural herita-ge, including the uniqueness of their wild pigs.

Babyrousa babyrussa: Vulnerable or Endangeredaccording to subspecies

Three extant subspecies of babirusa are curren-

tly recognized, all found in Indonesia:Babyrousa babyrussa celebensis from Sulawesi, B.b. togeanensis from the Togian Islands and B. b.babyrussa from the Sula Islands and Buru in theMoluccas. It has been extirpated from much ofSulawesi and is still threatened there andthroughout its range by hunting and habitatloss. The babirusa inhabits tropical rain forests,and in recent years large-scale commercial log-ging has posed a major and increasingly seriousthreat. They are one of the first animals tobecome locally extinct after logging or landopening. On Sulawesi 12,000 sq km of landhave been declared as wildlife protection areas,but as yet there are no national parks or otherwildlife reserves within the ranges of the othertwo subspecies. Effective conservation will require field statussurveys to develop management recommenda-tions for the enhanced protection of the spe-cies, particularly the least known but potential-ly most threatened races, B. b. togeanensis andB. b. babyrussa. The government should beassisted in its efforts to establish national parkson islands where these animals occur andrequire further protection. Hunting for subsi-stence and commercial purposes should beinvestigated with a view to its control or elimi-nation. Captive breeding programs should beinitiated for the two most threatened races,and fresh blood-stock should be introducedfrom the wild into the existing captive popula-tion of B. b. celebensis.

3. Non-native suids of conservation interest(Tab.2)Introduced and feral pigs should generally beregarded as exotic pests which should be con-trolled or eradicated wherever possible, howe-ver some populations warrant in-situ conserva-tion because they are representative of extinctor endangered taxa, are of anthropogenic orsocioeconomic significance, or are of uniquegenetic importance. There are three such suidtaxa in Southeast Asia.

Babyrousa b. babyrussa: Endangered

Although discussed above with the other racesof babirusa, this distinctively long haired formmay have been introduced to the Sula Islandsand Buru Island from southern Sulawesi whereit is now extinct. No matter what its origins,the race is severely threatened and in need ofconservation action.

Blouch R. A. Status of species, genetics and conservation - Communication

24 IBEX J.M.E. 3:1995

ex-Sus scrofa: Endangered - Andaman Islands Indeterminate - Nicobar Islands

These small pigs have descended from feralstock introduced at least 2,000 years ago, andhave evolved as an integral component of theirinsular ecosystems. Both the Andaman andNicobar populations were formerly assumed tobe endemic and are protected under Indianlaw. They are a primary food source for the iso-lated tribal societies inhabiting these islands,and may also have ritual and religious signifi-cance. A recent influx of immigrants has led tohigh levels of deforestation from logging, agri-cultural encroachment and other develop-ments. The pigs are also threatened by immi-grant poachers who use more efficient huntingtechniques than the tribals, including firearms,snares and dogs. A joint zoological-anthropolo-gical survey is needed to ascertain the presentdistribution, status and threats to the pigs, andto understand their role in the culture of theaboriginal tribes. Based on these findings,recommendations should be made for the pigs’enhanced future protection, taking intoaccount the legitimate rights and needs of theoriginal human inhabitants.

ex-Sus celebensis: Indeterminate

On Simeulue Island, northwest of Sumatra, thepresence of a highly modified form of S. cele-bensis gives clues to the origins of the island’shuman inhabitants whose language is most clo-sely related to Buginese or other southernSulawesi dialects. Thus apparently sometime inthe past, settlers arrived at Simeulue fromdistant Sulawesi, bringing with them the pigsthey had domesticated from the forests of theirhomeland. Other populations of wild pigs refer-

red to as feral S. celebensis are known fromseveral islands in the Moluccas and LesserSundas. Field studies and surveys are needed toclarify the distribution, status and systematicaffinities of these pigs. Anthropological com-ponents of such studies should investigate rela-tionships between the ethnic origins of localtribal groups and the distribution patterns ofwild pigs of varying derivation, as well as thecultural and socioeconomic importance ofthese animals. Protected areas need to be esta-blished, but management plans should alsoaddress the possibility that measures may needto be taken to control the population numbersof these feral pigs.

REFERENCESBRAATZ S., (1992) - Conserving Biological Diversity: A

Strategy for Protected Areas in the Asia-PacificRegion. Tech. Paper No. 193, Asia TechnicalDepartment Series. The World Bank,Washington, D.C.

GROVES C.P., (1981) - Ancestors for the Pigs: Taxonomyand Phylogeny of the Genus Sus. Tech. Bull. No. 3,Dept. of Prehistory, Australian National Univ.Press., Canberra, 96 pp.

GROVES C.P. & GRUBB P., (1993) - The EurasianSuids: Taxonomy and Description. In: Pigs,Peccaries and Hippos: Status Survey andConservation Action Plan. Oliver W.L.R. (ed),IUCN. Gland, Switzerland: 107-112.

OLIVER W.L.R. (ed.), (1993) - Pigs, Peccaries andHippos: Status Survey and Conservation ActionPlan. IUCN. Gland, Switzerland.

OLIVER W.L.R., COX C.R. & GROVES C.P., (1993) -The Philippine Warty Pigs (Sus philippensis and S.cebifrons). In: Pigs, Peccaries and Hippos: StatusSurvey and Conservation Action Plan. OliverW.L.R. (ed), IUCN. Gland, Switzerland: 145-155.

Status of species, genetics and conservation - Communication Blouch R. A.

25IBEX J.M.E. 3:1995

Taxon/Location/Status Remarks

Babyrousa b. babyrussa Otherwise extinct form probably introducedBuru and Sula Island from southern Sulawesi, threatened by loggingEndangered and settlers from Java; two reserves proposed

ex-Sus scrofa Feral, introduced 2,000 yrs ago; small (35-40 kg);Andaman Island-Endangered primary food for native tribes; threatened byNicobar Islands-Indeterminate deforestation and immigrant poachers

ex-Sus celebensis Feral on Simeulue, brought by settlers whoseSimeulue Island language related to Buginese; reserve proposed;Several island in Moluccas and Lesser Sundas little known of situation on other islands

Table 2: Non-native SE Asian Suids of Conservation Interest

26 IBEX J.M.E. 3:1995

1. Taxonomy and distributionFollowing Sanborn (1952), the wild pigs of thePhilippines have generally been attributed totwo, more widely distributed species, namely:the bearded pig, Sus barbatus, and the Sulawesiwarty pig, S. celebensis. Thus, the wild pigs ofthe west Philippine islands of Balabac, Palawanand the Calamian Group, which form part ofthe Sunda Shelf, are most closely related to thebearded pigs of Borneo, Sumatra and theMalaysian Peninsular, whilst those of the cen-tral (Visayas Islands) and eastern (Luzon,Mindanao and associated islands) Philippines,which form part of the Wallacean Region,were lumped with the Sulawesi pig.In a major review of the genus Sus, Groves(1981) confirmed the close relatedness of the

west Philippine pigs with S. barbatus, but reaf-firmed their separation as an (endemic) subspe-cies, S. b. ahoenobarbus. However, Groves alsoargued that the affinity of the central andeastern Philippine pigs with S. celebensis waspurely superficial and that these populationswere also more closely allied to S. barbatus; aview later endorsed by Mudar (1986). Groves(1981) also asserted that the central (Cebu andNegros) and eastern (Luzon, Mindoro,Mindanao and Jolo) Philippines populationswere not only distinct from those of thewestern Philippines, but were also distinct fromeach other. These regional populations weretherefore reassigned as two separated subspe-cies of S. barbatus, namely S. b. cebifrons and S.b. philippensis, respectively (Groves, 1981).

THE TAXONOMY, DISTRIBUTION AND STATUS OF PHILIPPINE WILDPIGS

Oliver W.L.R.IUCN/SSC Pigs and Peccaries Specialist Group. Park End, 28A Eaton Road, Norwich, Norfolk NR4 6PZ, U.K.

Keywords: Wild pigs, Suidae, Bearded pig, Warty pig, Sus barbatus, Sus celebensis, Sus philippensis, Asia, Karyotype.

IBEX J.M.E. 3:26-32

Abstract: Recent studies have revealed that there are three species and at least two subspecies of wild pigs in thePhilippines, of which two species and one subspecies are endemic.This is a larger number of endemic suid taxa thanany other country, with the exception of Indonesia. Within the country, the distribution of the native and endemicsuids follows broadly predictable lines, with divisions equating to the major “faunal regions” of late Pleistoceneislands. Thus the “warty” pigs east of Wallace’s Line in Luzon and Mindanao (including Samar, Leyte and, proba-bly, Bohol), i.e. S. philippensis, and those of the West-central Visayas Islands (Panay, Negros, Cebu and, probably,Masbate), i.e. S. cebifrons, are endemic at the species level; whereas those of Palawan and associated islands, i.e. S.b. ahoenobarbus, are closely related to the “bearded” pigs of the Sundaic Region (Borneo, Sumatra, MalaysianPeninsular, etc.) and are endemic only at the subspecies level. There is also evidence that the range of the nomina-te form of the bearded pig from Borneo, i.e. S. b. barbatus, extends to the small islands of Tawitawi and Sibutu inthe Sulu Archipelago. If this is the case, these are the only non-endemic wild pig populations in the Philippines.Unfortunately, however, the generally extreme levels of deforestation on most islands on which they occur, coupledwith intense hunting pressure, inadequate legal protection and the poor enforcement of existing legislation evenwithin most protected areas, have resulted in the systematic decline of all Philippine populations of these animals.These factors are especially apparent in the (west) Visayan region, where the endemic warty pig, S. cebifrons, isalready extinct or close to extinction on at least four (Masbate, Guimaras, Cebu and Sequijor) of the six islands inwhich it is known or believed to have occurred. It now survives only in a few small, isolated areas on Negros andPanay, where all remaining populations are declining as a result of continued habitat destruction and intense hun-ting pressure. These populations are also potentially seriously threatened by “genetic contamination” through inter-breeding with free-ranging domestic and feral pigs (unpubl. data). By comparison, S. philippensis remains relativelywidely distributed in most still-forested areas on the larger islands of Luzon, Samar, Leyte and Mindanao, where itoccurs in all of the principal national parks. It probably also still occurs on a number of the smaller islands withinthese regions, but is certainly threatened or extinct on some others. Further studies are likely to reveal genetic diffe-rences between some of the principal populations of this species, which is currently (but probably incorrectly)regarded as monotypic.

Status of species, genetics and conservation - Communication Oliver W.L.R.

27IBEX J.M.E. 3:1995

Even so, it was stressed that these were tentati-ve assignations owing to the dearth of museumspecimens from the Visayas Region (whereonly two skulls were available for examinationfrom Cebu, only one from Negros and nonefrom the other Visayan islands of Guimaras,Panay and Masbate) and the complete absenceof any comparative cytogenetic data, precludeda definitive assessment of the systematic rela-tionships of these populations.To a large extent this situation still obtains,though there have been some important deve-lopments in our understanding of the systema-tic relationships and genetic diversity of thePhilippine suids in recent years. These deve-lopments include the acquisition of a series ofskulls and mandibles from Negros (cebifrons)and Samar (philippensis) which, together withthe first photographs revealing the externalcharacters of the Visayan animals, not only ledGroves (1991) to reaffirm his assertion that thecentral and eastern pigs are more closely alliedto barbatus than to celebensis, but also tosuggest that these are sufficiently different frombarbatus and from each other to warrant sepa-ration as distinct species, namely S. cebifronsand S. philippensis, respectively (Groves, 1991;Oliver, 1991,1992). A description of thesesmall (S. cebifrons) to medium (S. philippensis)sized pigs is provided by Groves and Grubb(1993), who treat both species as monotypic,but acknowledge that S. philippensis appears tobe regionally variable in some characters andmay ultimately prove polytypic.The first studies of karyotypes and banding pat-terns of Philippine wild pigs have also yieldedimportant new information. In 1992, bloodsamples were collected from seven individualsof known origin (including two F1 captive-bred hybrids), representing five islands -Palawan, Culion, Mindoro, Luzon andMindanao - and the results were comparedwith those from similar studies of other speciesof Sus which have also been undertaken inrecent years. The diploid chromosome numberof the domestic pig and Asian and South-EastAsian populations of the Eurasian wild pig (Susscrofa) is invariably 38. The same number hasbeen found for S. barbatus, S. celebensis, S. ver-rucosus (the Javan warty pig) and S. salvanius(the pigmy hog). The preliminary results fromthe Philippine pigs are therefore of considera-ble interest. Of the seven pigs sampled, threepigs (a boar from Luzon and two sows fromMindanao) had 2n = 36 chromosomes, with acentric fusion between chromosomes 13 and

16 in the homozygous condition, and two pigs(both sows, one each from Culion andMindoro) showed 2n = 38 chromosomes, withchromosomes 13 and 16 separately present.This type of translocation is new, both to thedomestic pig and to the wild sp

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