THE CONTRIBUTION OF ANIMAL DOMESTICATION TO THE SPREADOF ZOONOSES: A CASE STUDY FROM THE SOUTHERN LEVANT

Horwitz L.K. * & Smith P. ***

Departme1lt of Evolution, Systematics and Ecology, The Hebrew University, Jerusalem, Israel.

* *Laboratory of Physical Anthropology, Department of Anatomy and Cell Biology, Hadassah Medical School, The Hebrew

U'1iversity, Jerusalem, Israel.

Abstract' - The advent of animal domestication was associated with marked changes in human-animalinteractions, a process that had a signiticant impact on both parties. In animals these changes are expres-sed in features such as biogeography, morphometry and pathology. In humans they arc reflected in chan-ges in health, disease and demography as well as in social complexity. Current research on the origin andspread of emerging diseases, suggests that many infectious diseases common today originated in the pro-cess of animal domestication. The archaeological record of the Southern Levant (Lebanon, SouthernSyria, Jordan, Israel and the Sinai Peninsula) provides an example of the possible role of animal dome-

I stication in spreading zoonoses.

Resume - La contribution de la domestication animale a la diffusion des zoonoses: Ie cas du Levantmeridional. Le debut de la domestication ani male a ete associe a des changements remarquables desinteractions homme-animal, ce qui cut d'importantes consequences autant sur les hommes, que sur lesanimaux. En ce qui concerne les animaux, ces changements ont eu des consequences de nature biogeo-graphique, morphometrique et pathologique. En ce qui concerne l'homme, ils peuvent ctre correles ades changements de l'etat de sante, de demographic et de complexite sociale. Des recherches en courssur l'origine et la diffusion des maladies, suggerent que plusieurs pathologies infectieuses aujourd'huifrequentes trouvent leur origine dans Ie processus de la domestication animale. La documentationarcheologique du Levant meridional (Liban, Syrie meridionale, Jordanie, Israel, Peninsule du Sinai")fournit des exemples du possible role joue par la domestication animale dans la diffusion des zoonoses.

\Key-words: Animal domestication, Zoonoses, Southern Levant.Mots des: Domestication animale, Zoonoses, Levant meridional.

1. IntroductionThe effect of the" Agricultural Revolution"on health status and nutrition has beendiscussed at great length in the scientific lite-rature (Cockburn, 1971; Cohen &Armelagos, 1984; Cohen, 1989). As notedby Brothwell (1978), it constituted" amixed blessing". On the one hand, it expan-ded the quantity and availability of food allyear round. On the other hand, it was asso-ciated with marked changes, often delete-rious, in the pattern of human disease.Several factors have been proposed toaccount for this shift in human health status.

IBEX J. MT. EcOL. 5:2000

IBEXJ. Mt. Ecol. 5: 77-84ANTHROPOZOOLOGICA31: 77-84

They include increased sedentism and popula-tion density, disease incidence and poorernutrition reflected in variety and quality offoods consumed (Cohen & Armelagos, 1984;Cohen, 1989; Smith & Horwitz, 1998).In this context, the role of animals as a sour-ce of infectious disease has been dealt with inthe zoological and archaeological literatureonly in a cursory manner (Brothwell, 1978;Cohen, 1989; Siegel, 1976). However, theimportant role of animals as hosts and car-riers of infectious diseases, at the presenttime (Twigg, 1980; Beran & Steele, 1994)suggests that changes in animal-human inte-

77

ractions resulting from the process of dome-stication, would have had a marked impacton the incidence and prevalence of infec-tious diseases in past communities.Evidence for this may be sought in theosteological remains of humans and theirdomesticates in archaeological contexts(Baker & Brothwell, 1980; Cockburn, 1971;Ortner & Putschar, 1985), as well as inongoing research into the origins and evolu-tion of pathogens (Greenblatt, 1998).

2. ZoonosesThere is considerable inter-specific variationin the susceptibility of animals to infectiousorganisms. In many cases; the animals act asa disease reservoir or intermediate hostwithout succumbing to the disease themsel-ves. Transmission to humans may be direct,for example, infection from saliva or bloodfrom rabid dogs, droplets in the air fromexhalation, sneezing or coughing or con-sumption of meat, blood or milk of a disea-sed animal (Cliff et at., 1998). It may also beindirect from accidental ingestion of infec-tious organisms from fecally contaminatedfood or drink, or transmitted by bites of avariety of insect vectors such as ticks, miteslice and fleas (Beran & Steele, 1994).Examples of some arthropod borne diseasesare Malaria and Dengue fever (mosquitoes),Lyme disease (ticks), typhus and plague(fleas). We propose that the short and inter-mittent contacts that took place betweenhumans and their prey prior to domestica-tion would have minimized opportunitiesfor infection by any means other than directconsumption of diseased animals. Followingdomestication contacts between humans andanimals increased with animals kept in closeproximity to human settlements. In somecases, they shared the same dwelling space,greatly increasing chances for disease tran-smission, and attracting commensals thatfurnished yet another source of infection.For humans, animal domestication was asso-ciated with:i. Close, direct and prolonged contact

78

between humans and animals including thekeeping of pets, feeding and stabling of ani-mals in or near human habitations. Milkingand sheep- shearing would have increasedthe intensity and duration of such contactwith animals and their by-products furtherfacilitating the transmission of diseases.ii. Increased sedentism, which is probably apre-condition for domestication, would haveincreased the risk of infection from poorsanitation. The associated increase in popu-lation size, which characterizes the Neolithicperiod, would have exacerbated the situationand contributed to the increased level ofinfection in these populations.iii. Use of animals for transport would havefacilitated greater human movement andexpand inter-group contact and trade ties.This would have increased the pool of peo-ple liable to infection.iiii. The keeping of herd animals imposesconstraints on settlement location-and espe-cially on the proximity to water. This wouldhave increased the likelihood of humaninfection from diseases such as malaria andschistosomiasis. ' \No systematic study has yet been carried outof pathologies associated with infectiousdiseases in faunal collections from differentperiods. However, many of the morphome-tric changes found in early herd animals(sheep, goat, cattle, and pig) appear to be atleast partially due to nutritional impoverish-ment, over-crowding and loss of mobilityfollowing domestication (Zohary et at. ,1998).Diseased animals which would have died inthe wild were perhaps able or even helped tosurvive in this new anthropogenic environ-ment, which coupled with the proximity ofother animals and humans would have hel-ped to maintain and spread diseases.Consequently, even the earliest domesticate,the dog, may have been the source of manyinfectious diseases. Most of the diseases tran-smitted by dogs leave no specific trace onthe bones or teeth. Some are fatal eventoday, others produce chronic ill health. Apartial list includes rabies and possibly

IBEX J. !vIT. ECOL. 5:2000

tHU"''';vi,,,,,,IIc~hm'ni,,,~ ,nd amo,bi,

~iseases such as Entamoeba hystolitica, which~auses cysts in the liver (Beran & Steele,

f994). Dogs also har~or ticks and fleas,while round worm, filanal worms and hyda-~id cysts are commonly transmitted to~lL1mansby accidental ingestion of fecallyIcontaminated food or water.piseases .commonly transmitt~d from her?animals Il1clude toxoplasmosIs, brucellosIsfrom sheep and goat milk, and tuberculosisfrom cattle. Both tuberculosis and brucello-sis may cause specific bony lesions (Beran &Steele, 1994). Cattle and, pigs also serve, asthe intermediate host of Cestodes, whichcause tapeworms in humans when the fleshfrom infected animals, is eaten.A variety of wild as well as domestic animalsmay be susceptible to tuberculosis (Beran &Steele, 1994). The disease may be air-borne,transmitted directly by saliva, milk or feces.Spores may also survive for a long time in thesoil. Anthrax is caused by Bacillus anthracis,which can survive for decades by becomingdormant and only becomes re-activatedwhen introduced into a conducive environ-ment inside an animal host. Spores in dirt ordust infect cUts in the skin or mucosa. Itaffects a wide range of species, both dome-stic and wild and today occurs worldwide(Beran & Steele, 1994). Humans are relati-vely more resistant to anthrax than cattle butprobably slightly less so than pigs and dogs.

3. A case study: the Southern Levant3.1. - AnimalsIn the Levant, plant and animal domestica-tion probably began in the Late Natufianperiod (10,500 bp) which witnessed theintensive eXploitation of cereals and pulses,as well as the first appearance of the dome-stic dog. However, the fidl spectrum of ani-mal domesticates is only completed by theend of the Iron Age (3,200 bp) (Tab. 1).Consequently, animal domestication in this

Iregion should be viewed as an ongoing pro-

I cess, which took place throughoUt much ofthe Holocene, rather than as single event.

IBEX J. MT. ECOL. 5:2000

Remains of the earliest domestic animal, thedog (Canis familiaris), have been identifiedfrom a number of Late Natufian (circa10,500 bp) sites in the southern Levant.They are often found in association withhuman burials suggesting that they mayhave been perceived as high status animals(Tchernov & Valla, 1997). Human diet atthis time was still based upon hunted andgathered f.1una, as well as intensive collec-tion of wild grasses and plants. This periodalso witnessed a marked increase in frequen-cies of commensal species such as the housemouse (Mus musculus), spiny mouse(Acomys cahirinus) and house sparrow(Passer domesticus) (Auffray et al., 1988;Tchernov, 1984) a factor that may be corre-lated with increased human sedentism andfood storage. Only by the late Pre-PotteryNeolithic B (PPNB) period (circa 8,500 bp),is there convincing evidence for the presen-ce of domestic goats (Capra hircus). This isshown by marked changes in skeletalmorphometry (high frequencies of twistedhorncores and a notable reduction in bodysize), as well as in age and sex profiles goats(Legge, 1996; Garrard et al., 1996).These features all point to a significant chan-ge in patterns of animal management andone, which differs markedly to that, associa-ted with hunting. At this time domesticsheep (Ovis aries) first appear in several sitesin the sOUthern Levant, all situated in, oradjacent to the Jordan Valley (Horwitz &Ducos, 1998). Sheep were probably intro-duced into this region from the north.By the late PPNB, species richness anddiversity in archaeological sites is reducedsuch that caprines are the main speciesexploited while other wild species such asgazelles, wild boar, aurochs, carnivores,birds and reptiles composed only a minorportion of the animal diet (Horwitz, 1996).Although there is no evidence for morpho-metric change in PPNB herd animals, manyof them show a relatively high incidence ofbone pathologies. Most common are exo-stoses (osteoarthritic deformities) primarily

79

SPECIES PERIOD WHEN DATE DOMESTICATED NUMBER OF SHARE[DOMESTICATED (bp uncalibrated) DISEASES

Camel Late Bronze Age 1,400 7Poultry Late Bronze Age II 1,400 26Rats and mice' Late Natufian 10,500 32Horse Chalcolithic 6,500 35Pig Pottery Neolithic 7,500 42Sheep and goats Late PPNB 8,500 46Cattle Pre-Pottery

Neolithic C 8,100 50Dogs Late Natufian 10,500 65

,Commensals

Tab. 1- Number of diseases shared between domestic animals and humans (tram: McNeill, 1976)

found on goat phalanges probably the resultof reduced mobility (Kohler-Rollefson,1997; Goring-Morris et ai.) 1995). They arealso found in PPNB cattle from Jericho and'Ain Ghazal (Clutton-Brock, 1979; Kohler-Rollcfson, 1986). This contrasts to the situa-tion for gazelle from the same sites whichshow no pathologies. One goat from the siteof 'Ain Ghazal had a healed fracture, indica-ting that the animal was kept and cared forfollowing this trauma. Another caprine fromthe same site exhibited resorption of bone inthe limb extremities. This was interpreted asevidence for intentional tethering (Kohler-Rollefson, 1997). Although these patholo-gies are not associated with specific diseases,they do provide some indication for stress,presumably due to the conditions in which-these early domestic animals were kept.Morphometric change in herd animals attri-buted to domestication, is observed by thePre-Pottery Neolithic C (circa 8,100 bp)(Ducos & Horwitz, 1998). By the PotteryNeolithic (7,500 bp) the faunal assemblagesin the southern Levant are dominated bythe main Near Eastern domestic animals:sheep, goats, cattle and pigs. Domesticdonkeys first appear in the archaeozoologi-cal record of the southern Levant in thisperiod. The donkey appears to have been

80

introduced as a domesticate into this regionfrom the south, possibly Egypt. Based onclay figurines from the Pottery Neolithic andthe subsequent Chalco lithic period, theyappear to have served primarily as pack ani-mals (Ovadia, 1992). Domestic horses mayhave been present in the southern Levant bythe Chalcolithic period (Grigson, 1993).However, their remains do not occur in anyfrequency prior to the Iron Age. It is accep-ted that horses were introduced as a dome-sticate into the southern Levant, either fromEgypt or the Northern Levant. Camels tooare clearly an introduced species. There isambiguous data concerning their presence inthe region in sites dating to the MiddleBronze Age, but by the Late Bronze Agethey are definitely present. Poultry arefound in substantial numbers by the IronAge, but may have been introduced in theLate Bronze Age. Subsequent additions tothe domestic fauna of the Southern Levantinclude the cat, domestic goose, pigeon andwater buffalo introduced into this regionmore recently. The first evidence of milkingdates to the Chalcolithic period, where cera-mic vessels interpreted as milk churns, arecommon (Amiran, 1969). Cortical bonethickness in sheep and goats shows a markeddecrease from Chalcolithic to the Early

IBEX J. MT. ECOL. 5:2000

,ronze Age, and remains low in laterbriods. This has been attributed to negative~lcium balance resulting from the combi-~d effects of poor nutrition and intensive~ilking. It indicates that only in the Early4lronze Age were herds of sheep and goat~ploited on a large scale for their milkSmith & Horwitz, 1984).4~.2. HumansChronic diseases are associated with impai-Ijed health and growth and frequently result.n early death. Identification of a specific4jisease is however possible only if it produ-~es a characteristic bony lesion that can be~ecognized in the teeth or bones.4Many diseases when first introduced into ailllman community are fatal, such that mor-tality may result rapidly before any signs arefound in the skeletal record. Evidence forthis may be indirect such as abandonment of'sites or mass burials, where the age profile-differs from that expected through naturalmortality. The number of skeletal remainsrecovered from different periods is too smallto attempt a detailed reconstruction ofpaleodemography. However, it does providean opportunity to assess changes in life-spanfrom an examination of the age at death.These data are calculated relative to the pro-portion of total adults identified, excludingchildren as their remains are poorly preser-ved or may have been buried outside ofcemeteries. In the Natufian and Neolithicperiods, only some 10% of adults appear tohave survived beyond the age of 50 years.This frequency drops in the Chalcolithicperiod, where no individual appears to havesurvived to the age of 50. By the MiddleBronze Age (circa 3,750 bp) there is consi-derable amelioration with between 20-30%surviving to 50 or more years of age. In theIron Age, the situation deteriorates slightlywith less than 20% survival beyond the ageof 50 years (Smith & Horwitz, 1998).

I

There is good evidence for increased environ-mental stress in Neolithic and Chalcolitl1icpopulations. This is shown by the high fre-quency of hypoplastic defects in the teeth,

growth arrest lines in the long bones andosteoporosis in young adult females. Inaddition, cribra orbitalia and parietal pitting,show a concomitant increase (Smith et al.)1984; Smith & Horwitz, 1998). These con-ditions are indicative of iron deficiencyeither through dietary deficiency, chronicanemia associated with intestinal infectionsor genetic factors affecting red blood cellformation (Stuart-Macadam & Kent, 1992).The first definite evidence of a specific disea-se transmitted by animals is found in themid/late PPNB levels at the site of 'AinGhazal, where two individuals have beendiagnosed as suffering from tuberculosis(El Najjar et aI., 1997). A number of otherspecimens from the late PPNB site of Bastaand the PPNC of Ain Ghazal, also showbony changes that may be attributed totuberculosis (Schultz & Scherer, 1991).In the Chalcolithic period similar pathologi-cal symptoms associated with tuberculosis,but not pathognomic of the disease, havebeen identified from several sites includingShiqmim, Kissufim and Gilat (Zagerson &Smith, in press). The Early Bronze Age siteof Bab edh-Dhra also shows evidence oftuberculosis (Ortner, 1979).

4. DiscussionThe data presented here show that there is aclear association between animal and plantdomestication and deterioration in health ofhumans and their herds. It seems that therewas a significant deterioration in the healthstatus of early Neolithic populations relativeto that of the Natufian hunters and gatherersthat preceded them. This trendbegins befo-re the domestication of herd animals in theearly phases of agriculture (Pre- PotteryNeolithic A period). However, it should beremembered that dogs were already dome-sticated by this time and are a major sourceof zoonoses. Deterioration in health isreflected in shortened life expectancy, theincreased incidence of developmental defectsand in osteoporosis and cribra orbitalia.All these conditions may be associated with

81

impoverished diet and/or failure to absorbfoods (Smith et al., 1984). We cannot atthis stage assess the relative role played bychanging diet versus disease on humanhealth. However epidemiological studies ofThird World populations have found thatdisease rather than diet is the main factorresponsible for chronic ill-health (Cliff et al.,1998). In the Near East the process of ani-mal domestication occurred over severalmillennia. In terms of zoonoses, this wouldhave meant the constant introduction ofpotentially new sources of disease as diffe-rent animals were adopted into the anthro-pogenic domain. This scenario differs fromthat occurring in regions such as WesternEurope, where domestic animals were intro-duced as a package within a relatively shortperiod oftime. We might then expect to finda less severe, but more prolonged impact onNear Eastern populations, where new disea-ses were introduced over a long period.The earliest well documented evidence forearly transference of disease from animals tohumans is the presence of tuberculosis inhumans in the late PPNB, populations thesame time as a high frequency of foot patho-logies is found in herd animals.The pathology in herd animals provides con-vincing evidence for reduced mobility inthese animals and probably reflects theirdeleterious living conditions. Such circum-stances are known to be excellent breedingplaces for zoonoses such as bovine tubercu-losis and provide a basis for the transmissionof zoonoses in this period. The majority ofdomestic animals found in the SouthernLevant (perhaps with the exception of thedog and goat) appear to have been introdu-ced into the region as domesticates and did notundergo autochthonous domestication here.Consequently, it is expected that these ani-mals would have brought with them newdiseases for which neither the local wildfauna nor human communities would havedeveloped any immunity. Although there isno direct for diseases introduced by theseimported species, the continued deteriora-

82

tion in human health until the MiddlelBronze Age period suggests that the intro-duction of new species was associated withnew health challenges. The data are stillincomplete, and we have highlighted hereonly one facet of the many changes inhuman society associated with animal dome-stication. A further indication of the contri-bution of human-animal contacts to thespread of diseases is shown by the correla-tion present between the time of domestica-tion and the number of diseases shared withhumans (Tab. 1). Thus, the species mostrecently domesticated, poultry and camels,share fewest diseases with humans while theearliest domestic animal, the dog, showsmost (McNeil, 1976). These data plainlyillustrate the direct connection between theprocess of animal domestication and theincrease in infectious diseases in humans.They provide a starting point for integratingdata from molecular biology with that deri-ved from the archaeological record.

REFERENCES \

AMlRAN, R. (1969) - Ancient Pottery of theHoly Land. Jerusalem, Masada Press.

AUFFRAY,J-c., TCHERNOV, E. & NEVO, A.(1988). Origine du commensalisme de lasouris domestique (Mus musculus domesticus)vis-a-vis de l'homme. Cahiers de RechercheScientifique de Paris 307: 517-522.

BAKER, J, & BROTHWELL, D. R. (1980) -Animal Diseases in Archaeology. London,Academic Press.

BERAN, G.w. & STEELE, J,H. (1994) -Handbook ofZoonoses. 2nd edition. SectionA: Bacterial, Rickettsial, Chlamydial, andMycotic. Boca Raton, CRC Press.

BROTHWELL, D.R. (1978) - The question ofpollution in earlier and less developedsocieties. In: Logan M.H. & Hunt E.E.(Eds.), Health and The Human

Condition. Massachusetts, DuxburyPress.: 129-136.

CUFF, A., HAGGETT, P.RAYNOR, M. (1998)

& SMALLMAN-

- Deciphering

IBEX J. MT. ECOL. 5:2000

Global Epidemics. Cambridge Studies inHistorical Geography 26. CambridgeUniversity Press.

CLUTION-BROCK,J, (1979) - The mamma-lian remains from the Jericho Tell.Proceedings of the Prehistoric Society 45:135-157.

COCKBURN,T.A. (1971) - Infectious Diseasesin Ancient Populations. CurrentAnthropology 12: 45-62.

COHEN, M.N. (1989) - Health and the RiseofCiJlilization. Yale University Press.

COHEN, M.N. & ARMELAGOS, G.J.Eds.(1984) - Paleopathology at theOrigins of Agriculture. Academic Press.

Ducos, P. & HORWITZ,L.K. (1998) - Theint1uence of climate on artiodactyl sizeduring the Late Pleistocene-EarlyHolocene of the southern Levant.Pa/Corient 23: 229-247.

EL-NAJJAR,M. , AL-SHIYAB,A. & AL-SARIE,1. (1997) - Cases of tuberculosis at 'AinGhazal, Jordan. Pa/Corient 22: 123-128.

GARRARD,A.N., COLLEDGE,S., & MARTIN,L. (1996) - The emergence of cultivationand pastoralism in the marginal zone ofthe Levant. In: HARRIS D.R. (Ed.) TheOrigins and Spread of Agriculture andPastoralism in Eurasia. London:University College Press.: 204-226.

GORING-MoRIus, N., GOREN, Y., HORWITZ,L.K., BAR-YOSEF,D. & HERSHKOvrrz,1.(1995) - Investigations at an earlyNeolithic settlement in the LowerGalilee: Results of the 1991 season atKefar Hahoresh. Atiqot) 27: 37-62.

GREENBLATI,C. L. (1998) - Digging forPathogens. Rehovot Balaban Publishers.

GRIGSON,C. (1993) - The earliest domestichorses in the Levant? New finds from theFourth millennium of the Negev. Journalof Archaeological Science) 20: 645-655.

HORWrI"Z,L.K. (1996) - The impact of ani-mal domestication on species richness: Apilot study from the Neolithic of thesouthern Levant. ArchaeoZ(}ologia)8: 53-70.

HORwm, L.K. & Ducos, P. (1998) - Aninvestigation into the origins of domestic

IREX L MT. EcoL. 5:2000

sheep in the southern Levant. In:Buitenhuis H., Bartosiewicz L. & ChoykeA.M. Vill. III. (Eds.) Archaeozoology of theNear East) Groningen, ARC Publicaties.pp. 80-94.

KOHLER-RoLLEFSON, 1. (1986) - Bonepathology as indication of domestication.In: Cultural Attitudes to Anima/J'Including Birds) Fish and InvertebratesVol. 1, Section on IIEarly Husbandryand the Exploitation of domestic Stock ".The One World Archaeological Congress,Southampton, Allen & Unwin.

KOHLER-ROLLEFSON,1. (1997) - Proto-ele-vage, pathologies and pastoralism: Apost-mortem of the process of goatdomestication. In Gebel, H-G., Kafafi, Z.& Rollefson, G.O. (Eds.), The Prehistory ofJordan II. Perspectivesfrom 1997. Berlin, exOriente. pp. 557-565.

LEGGE,A. (1996) - The beginning of capri-ne domestication in Southwest Asia. InHarris D.R. (Ed.): The Origins andSpread of Agriculture and Pastoralism inEurasia. London, University CollegePress. pp. 238-262.

MCNEIL,W.H. (1976) - Plagues and People.New York, Anchor Press.

ORTNER,D.J, (1979) - Disease and morta-lity in the Early Bronze Age people of Babedh-Dhra, Jordan. American Journal ofPhysicalAnthropology) 51: 589-597.

ORTNER, D.J. & PUTSCHAR,w.G. (1985) -Identification of Pathological Conditions inHuman Skeletal Remains. WashingtonD.C., Smithsonian Institution Press.

OVADIA,E. (1992) - The domestication of theass and pack transport by animals: A case oftechnological change. In: Bar-Yosef O. &Khazanov A. (Eds.), Pastoralism in theLevant. Madison Wisconsin, PrehistoryPress. Pp. 19-28.

SCHULZ, M. & SCHERER, A. ( 1991 ) -Human skeletal remains. In: Nissen H.J"Muhsein M. & Gebel H-G. (Eds.),Report on the Excavations at Basta.Annals of the Department of Antiquitiesof Jordan 35, 95-98.

83

SIEGEL, J. (1976) - Animal palcopathology:Possibilities and Problems. Journal ofArchaeological Science 3, 349-384.

SMITH, P. & HORWITZ, L.K. (1984) -Radiographic evidence for changing patternsof animal exploitation in the southernLevant. Journal of Archaeological Science,11: 467-475.

SMITH, P. & HORWITZ, L.K. (1998) -Culture, environmeand disease: Paleo-anthropological findings for the southernLevant. In: Greenblatt C.L. (Ed.)Digging for Pathogens. Rehovot, BalabanPublishers. pp. 201-239.

SMITH, P., BAR-YOSEF, O. & SILLEN, A.(1984) - Archaeological and skeletal evi-dence for dietary change during the LatePleistocene/Early Holocene.in the Levant.In: Cohen M.N. & Armelagos G.}. (Eds.)Paleopathology at the Origins of Agriculture.

Academic Press. pp. 103-136.

84

STUART-MACADAM,P. & KENT, S. Eds.( 1992) - Diet, Demography and Disease:Changing Perspectives in Anaemia. NewYork, Aldine de Gruyter.

TCHERNOV,E. & VALLA,F. (1997) - Twonew dogs and other Natufian dogs, fromthe southern Levant. Journal ofArchaeological Science, 24: 65-95.

TWIGG,G.!. (1980) - A review of the occur-rence in British mammals of the majororganisms of zoonotic importance.Mammal Review, 10: 139-149.

ZAGERSON,T. & SMITH, P. (in press) -Chapter 10. The Human Remains fromKissufim Road. In: Y. Goren & P. Fabian(Eds.) Kissufim Road Final Report,Israel Antiquities Reports.

ZOHARY,D., TCHERNOV,E. & HORWITZ,L.K. (1998) - The role of unconsciousselection in the domestication of sheepand goats. Journal of Zoology London.

IBEX J. MT. ECOL. 5:2000