Two New Dogs, and Other Natufian Dogs, from the Southern Levant

Journal of Archaeological Science (1997) 24, 65–95

Two New Dogs, and Other Natufian Dogs, from the Southern Levant Eitan Tchernov The Department of Evolution, Systematics and Ecology, The Hebrew University of Jerusalem, Jerusalem 91904, Israel

François F. Valla Laboratoire d’Ethnologie Préhistorique, 44, Rue de l’Amiral-Mouchez, 75014 Paris, France (Received 31 May 1995, revised manuscript accepted 10 January 1996) Two new fairly complete remains of dogs were uncovered from a burial in Hayonim Terrace (northern Israel), dated to the first half of the 11th millenium () (Late Natufian). This burial contained the remains of three humans associated with an elaborate construction. A detailed analysis of these dogs, and a comparison with all known Natufian remains, suggested that genuine dogs were already living around and within human habitations during this period. While most studies on early dogs were carried out on post-Natufian material, a period during which intentional selection could have already been widely experienced, we show that the evolvement of Natufian dogs were the product of unconscious selection of commensal wolves quasi-isolated under the special anthopogenic habitats created within and around Natufian sites, and at least ritually assimilated to human society. There is no evidence that Neolithic dogs are direct descendants of Natufian ancestors. Their multiregional origination is a widely accepted phenomenon. We suggest that Neolithic dogs were either domesticated anew, or were introduced from elsewhere to the southern Levant. Although the number of specimens of Natufian dogs is still small, evidently the emergence of some patterns clearly indicate that all them were already markedly different from the recent southern Levantine wolves and, in spite of their widely ranged morphology, constituted a unique group. It is shown that the shortening of the muzzle mainly affected the anterior part of the snout, while the posterior region remained practically unchanged. In this respect they seem to display a typical case of paedomorphosis. The simultaneous diminution of the carnassials, and other teeth, alongside with the snout, was marked enough that no crowding of the teeth took place by any of the Natufian dogs. The disproportional reduction of the snout versus teeth that caused the ‘‘crowding’’ phenomenon is only known from later periods. ? 1997 Academic Press Limited

Keywords: DOG, WOLF, NATUFIAN, SOUTHERN LEVANT, SEDENTISM, COMMENSALISM, DOMESTICATION, HAYONIM TERRACE.

animals, dogs were somehow already assimilated to human society. If this was the case then the practice of burying dogs with people is a strong indicator, independent of biological factors, which through time induced morphological changes. According with Benecke (1987) the earliest definite record (a single jaw fragment) of domesticated dog came from Bonn-Oberkassel in central Europe, found in a grave associated with an old man and a young woman (Nobis, 1981), dated to the Magdalenian period, c. 14,000 , but defined by Benecke as an ‘‘early Mesolithic dog group’’ (p. 49). Accepting this isolated find shows yet again that human–wolf interaction with animals came about independently in many places, but mostly among sedentary human populations with long-term interactions with animals. Skeletal remains reported as those of early dogs derived from early archeological contexts (dating from approximately 9000–14,000 years ago) are scattered in

Introduction articular interest has always been attached to the origins and early history of domestic dogs. Bearing in mind the vast number of publications concerning the emergence and evolution of dogs, we would not dare produce yet another if there were not new interesting evidence, in this case early dogs found in a very special anthropogenic context, primeval which may shed new light on the problems of their earliest origins. Human–dog burials are well known from the Neolithic onward in many parts of the world (see for examples Bökönyi, 1973, 1974, 1983; Lechevallier et al., 1982; Bonnet et al., 1989). If our analysis of the Natufian evidence is correct, the appearance of such burials is linked to ritual activities which, in some way, refer to the reality of everyday village life, where people and dogs co-existed from birth to death. As commensal

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different regions, from the Middle East (Davis & Valla, 1978; Turnbull & Reed, 1974) to central and northern Europe (Benecke, 1987; Degerbol, 1961; Morey, 1992; Nobis, 1981) and into North America (Grayson, 1984). But we agree with Olsen (1985) that until now the earliest known domesticated wolves ‘‘. . . could not be stated with certainty, thus most of the early dates for dog domestication must be questioned and set aside until material complete enough . . .’’ (p. xii). Yet the Natufian dogs found in the southern Levant (Davis & Valla, 1978; Dayan, 1994) are exceptional in having not only unique morphological size and proportions compared with the local population of recent wolves, which definitely distinguishes them as a different taxonomic entity, but show a direct yet complicated cultural association with humans, being intentionally placed in an intimate position in common burials. In spite of Olsen’s (1985) criticism of the validity of the identification of isolated finds of ‘‘small wolves’’ as evidence for early domestication of dogs (Turnbull & Reed 1974; Davis & Valla, 1978), it now seems, following the discovery and analysis of more fossil remains from Natufian levels, that genuine dogs were already living around human habitations during this period. We would also not dare to come out with yet another publication on eastern Mediterranean early dogs soon after Dayan’s (1994) enlightening paper, if we did not have some new material and different views concerning their earliest beginnings in the southern Levant. In this paper we also tend to expose in more details the raw data of all other Natufian dogs in comparison with the recent southern Levantine wolves which will enable a basis for further comparison with the much more widespread Neolithic dogs. It has been frequently claimed (Zeuner, 1963; Lawrence, 1967; Clutton-Brock, 1970, 1981; Epstein, 1971; Bökönyi, 1973, 1974, 1983; Olsen, 1985) that a key to a clearer understanding of canid domestication lies with the very morphological changes that provide the basis for identification of early dogs. These changes include overall size reduction, shortening of the facial region of the cranium and, in the earliest specimens, large crowded teeth. It has also been frequently observed that changes in cranial morphology appear to reflect paedomorphosis, or the retention of juvenile characters into adulthood (Weidenreich, 1941; Zeuner, 1963; Epstein, 1971; Clutton-Brock, 1981, 1984; Morey, 1992; Wayne, 1986a,b,c). Yet we have to take into account that most of these studies were carried out on post-Natufian dogs, when intentional selection was the rule. We do not believe that the creation of Natufian dogs was either intentional or mentally preconceived. We have no solid evidence on which to base the notion that southwest Asian Neolithic dogs are direct descendants of the Natufian line. It is possible that Neolithic dogs were either domesticated anew in this region, or were introduced from elsewhere. The multiregional origination of domestic dogs is a well-known

phenomenon. In this paper we will avoid any discussion concerning Neolithic dogs, even those from the southern Levant.

The Archaeology of the Two New Dogs from Hayonim Terrace Hayonim cave and its terrace in front is situated at the western edge of upper Galilee (Figure 1). The cave is known to have been intermittently occupied from the early Middle Paleolithic onward. A long sequence deposits running from early Mousterian, through Levantine Aurignacian, Kebaran and Natufian to Historic periods has been ascribed Bar-Yosef & Tchernov, 1966; Bar-Yosef, 1983; Belfer-Cohen, 1988; Bar-Yosef & Belfer-Cohen, 1989) from the cave. The Geometric Kebaran, Natufian and final Neolithic deposits on the terrace are not stratigraphically connected with those in the cave. Along with other Natufian structures, two curvilinear constructions accompanied by seven graves have been identified as semi-subterranean dwellings (Valla et al., 1988, 1991). One of these graves is of particular interest since it contains the remains of three humans (Homo 7, 8 and 10) and two dogs. Twelve 14C dates have been obtained, and have shown a range from 16,810&210  (OxA 2974) to 6970&80  (OxA 1900). Five of them fall within the 12 millennium  and indicate an Early Natufian or an early Late Natufian period. However the analysis of the material suggests a Late Natufian age. Most lunates are made with bipolar retouch, the bulk of the bone tools are points, the skeletons in the graves are not decorated, art is present but very unusual. For those reasons we feel confident in assigning a Late Natufian age to the occupation of the terrace, placing it mainly within the first half of the 11 millennium . Moreover, the grave with the dogs was probably dug after the abandonment of the nearby house (structure 9) and hence does not belong to the earliest Natufian settlement at the place. For comparison, the dog remains from Kebara cave (Figure 1) can be assigned to an early phase (c. 12,500–12,000 ) and those from Eynan (=Mallaha) to a late phase (c. 11,500 ) of the Early Natufian, while the Shukhba dogs belong to the Late Natufian. The finds from el Wad could be either Early or Late Natufian since the exact layer could not be defined. The sequential events of the burials of Homo 7, 8 and 10 with the dogs can be interpreted as follows: an egg-shaped pit was excavated into which the bodies of two dogs were placed. Dog 1 was lying on its right side in an elongated position (Figure 2). Its head was raised along the neck, following the curvature of the grave. The forelegs were folded one on top of the other. The hind legs were extended and went up the edge of the pit, the left foot approached the skull of Homo 10 (Figures 3 & 4). The muzzle of dog 2 was near the

Dogs from the Southern Levant 67

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km Figure 1. A map (258,6 N; 170,6 E; Alt. 250 m) showing all the Natufian sites where dogs were recorded and studied, and other principal Natufian sites to illustrate the general distribution of the Natufian settlements in the southern Levant. -: Natufian sites with dog remains; ,: other main Natufian sites.

shoulder blades of dog 1 and its head was resting on the lower jaw. The exact position of the body of dog 2 is difficult to ascertain since some bones are missing (the upper part of the vertebral column, some ribs and the shoulder blades; some of the limb bones were incomplete. The lower part of the axial skeleton lies on top of the left femur, its proximal end pointing toward the muzzle. The animal may have rested on its chest, while the forelegs on each side of the body which may have been curved around a block of limestone. A flat block of limestone was laid on top of the chest of dog 1 and skull of dog 2. Under that stone the vertebral column of dog 1 was separated in two parts (Figure 5). Two tortoise shells of Testudo graeca

(Testudinidae, Chelonia) were found, one near the right elbow of dog 2; the other near the lower part of the vertebral column of dog 1. The body of Homo 8 (probably a male) was deposited on top of the dogs, reclining on its right side in a contracted position. His head lay on the slightly sloping bottom of the pit near the forehead of dog 1 and partly above the tortoise shells near the elbow of dog 2. The skull was laid between the frontal bone near that of the dog and the occipital bone close to the tortoise shells. The head is bent towards the right shoulder, the face looking toward the knees in the center of the grave. The pelvis is on its side on top of the back legs of dog 1 (right tibia and left femur). The

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Figure 2. Map of grave H.7-8-10 at Ghayonim Terrace. This grave was found among other Natufian structures: two houses; a pit, possibly for storage; six other burial pits, etc. The body of three individuals and two dogs were deposited in two layers. At the lower part (right) two individuals and the dogs were identified. The third individual (left) was found above. The grave was closed by a cover of cobbles not seen on this figure. Note the grid in order to restitude the position of H.7 on top of H.8.

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68 E. Tchernov and F. F. Valla

Dogs from the Southern Levant 69

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Figure 3. Vertical photograph of the dog’s remains after removing H.8. The window at the top right shows the relative position of two animals: the bones of dog 1 appears in black. The part of the skeleton of dog 2 (white), which was under the chest of H.8, is missing as were the bones of H.8 himself in the area.

right knee makes an angle of about 25); the left one is folded on top of it at about 40). The right tibia is sloping up on the edge of the grave so that the base of the foot is about 10 to 15 cm higher than the base of the distal epiphysis of the femur, and almost 20 cm higher than the highest part of the skull. The angle made by the right tibia and foot is c. 110); that made by their left counterpart is c. 90). Both feet converge toward the toes (Figure 2). The right arm is folded at c. 90). The wrist is in between the knees. The hand is in the prolongation of the forearm with the second and third phalanges folded. Metacarpals and phalanges are seen laterally, one on top of the other. Nothing was found of the left arm and only a few bones of the hand suggest that it was folded on the left side of the body, making an acute angle at the elbow. The right humerus lies across the neck of dog 2, the elbow being on top of the lower part of the back-bone of the same animal

while the ulna and radius cross the lower part of the backbone of dog 1. Homo 10 is linked to this stage of the grave by the left leg of dog 1 which is resting on its skull as already noted. Only a few bones of this individual are preserved. The sagital part of the skull is about 10 cm behind the pelvis and 30 cm behind the heels of Homo 8. The top of the bone lies at the same altitude than the top of the left femur and calcaneum of Homo 8. The face has disappeared; it was probably looking in the opposite direction to the face of Homo 8. Only a fragment of the lower jaw was found in between the bones of the arm. The postcranial elements are restricted to some cervical vertebras, the left clavicle and shoulder-blade are still connected, the bones of the left arm are in a poor state of preservation and a very limited number of bones from the hand were found. Broadly speaking, Homo 10 seems to be resting on a more or less flat surface, but the left leg of

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10 cm

Figure 4. Vertical photograph of H.8 and H.10. H.8 has a large stone on his chest. The stone standing in front of his face bears a cup mark. The head of dog 1 was found near the skull of H.8 and one of his hind legs was on top of the skull of H.10. The skull of dog 2 can be seen behind the shoulderblade of dog 1. Note the tortoise shell behind the skull of H.8.

dog 1 slopes down 17 cm from the phalanges which rest on the skull of Homo 10 to the distal epiphysis of the femur under the pelvis of Homo 8 (Figures 2, 3 & 4). At this stage some limestone blocks of were deposited in the grave. The largest one (25#15#15 cm) was put on the chest of Homo 8. Two other blocks, one flat (16#15#6 cm), the other rounded, occupied the space limited by the angle made by the right arm of Homo 8. Worth noting are also the blocks laid on top of the head of Homo 8 and Homo 10, and a flat stone along the left tibia of Homo 8, covering its right hand. Obviously, the stones were deliberately deposited, since they are in direct contact with the bones (with the

exception of the block on the chest of Homo 8 under which there are no bones) (Figures 2, 3 & 4). After the bodies (at least of Homo 8 and the dogs) had been covered with soil a third individual, Homo 7, was placed in the grave. He was deposited on his chest, so that his right shoulder lay on top of the skull of Homo 8. The skull lay on its right side, the face looking north-eastward towards the cliff. The scapular belt is bent in such a way that the left clavicle abuts the chin. Both humeri are diverging from the chest, the left forearm and hand are missing. The right elbow makes an angle of about 90), pointing the edge of the big stone on the chest of Homo 8. The wrist is at 90) to the forearm.

Dogs from the Southern Levant 71

10 cm Figure 5. Vertical photograph of H.7. This individual is lying on his chest. A large stone rests on the body. Two horn-cores of a gazelle are seen under the right arm. Another gazelle’s horn-core was found leaning against a stone on top of the right shoulder. The lower part of the skeleton is missing.

The chest of Homo 7 is preserved but the bones are badly crushed by a heavy block of limestone (22#16#19 cm) resting on them. A smaller block may have been purposefully set on the skull which is deformed in the area of the occipital bone (Figures 2 & 5). Yet the intentional deposition of gazelle (Gazella gazella) horn cores with the body of Homo 7 is in no doubt of intentional burial of an animal in this burial. A large horn core of a male was found on the top of the right shoulder in contact of the large stone on the chest. Two other horn cores and their supporting frontals were laid under the chest and the right arm. They lay approximately parallel to the forearm, both points of the horn cores passing under the humerus. The lower part of the body of Homo 7 disappeared. The grave was probably closed after Homo 7 had been deposited. There are some indications of a post-

depositional erosion event which may be responsible for the disappearance of the bones missing from Homo 7. Following this event the grave was covered with a protecting layer of stones. Later disturbances may have occurred when a fire pit was dug nearby. At this stage of the study we may assume that no further burial activity took place in the grave. The humans, animals and stones were rapidly buried. Since there is no clear indication to the contrary we conclude that the human (with possible exception of Homo 10) and dogs were introduced as complete corpses without any dismembering. The humans were probably buried with clothes which, combined with the stones, may have created free spaces, facilitating uneven decomposition and allowing some bones to move. As it stands, grave H.7–8–10 is one of the most complex sepultures known in the Natufian culture.

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Some graves have more individuals in them, a few others displayed animal remains (one grave from Eynan (=Mallaha) revealed a puppy; another one— again from Mallaha—exposed some horn cores of gazelles; a tortoise shell is known in a grave from the site of el-Wad) and many contain stones (Bate, 1937 Stekelis & Yizraely, 1963; Valla, 1975, 1977; Perrot et al., 1988). None of them included all these together. It cannot be by chance that dog 1 has his head near the skull of Homo 8 and a foot resting on the skull of Homo 10. Similarly the big stones on the chests of Homo 8 and 7, and the smaller ones on each skull Homo 8 and 10 (and possibly also Homo 7) were deliberately set there. The same is obvious for the stone with a cup-mark standing near the head of H.8. Finally the dualism: two humans, two dogs, two tortoise shells, at the lower level of the grave, as well as the opposition–human–dog–tortoise–stones (one with cup-mark) at that level; while human–gazelle–stone at the upper level. All this appear as an elaborate construction which can safely be interpreted as part of a rite. Another manifestation of what seems to be the same myth on a floor in a house at Eynan (=Mallaha) (Valla, 1988). There, within a distance of about 1 m, a human cranial cap was associated with a dog’s half jaw and the frontal bone of a gazelle with its horn cores attached. The interest of this document is to provide an example where three of the main elements combined in the graves intervene. But in this context each of them appears as a selected bone. This throw into special relief on their different treatment in the graves where gazelles still show up as a selected bone whereas human beings and dogs appear as complete bodies. In other words, we observe a strong contrast between the way gazelles are treated, i.e. never buried, and the treatment of humans and dogs (sometimes buried, sometimes not). Moreover the representations of humans and dogs both change together. In all likelihood this is a translation by the ritual of the status respectively achieved in the society by the two animals. Gazelles are obviously linked to human beings not only because they were heavily exploited as the dominant diet of the Natufian people, but because they were culturally controlled (Horwitz et al., 1991; Cope-Myashiro, 1992; Tchernov, 1993, yet never found in human burials. For dogs, on the other hand, there is no evidence for their being eaten. Living within or around villages their bodies were apparently removed from the living area, but probably under special circumstances they have been buried with people.

Earlier studies of Natufian Dogs in the southern Levant Dayan (1994) has already discussed in some details the earlier studies of Natufian Dogs in the southern Levant. We will only show that at present, when more

material came to light, the differences between the local wolves and the Kebaran wolves as was clearly shown by Dayan (Davis, 1981; Dayan et al., 1992), and the early Natufian dogs are conspicuous, and that the Natufian population represents a unique group, showing many traits which are utterly different from either Kebaran or recent wild wolves. Moreover, we tend to show that not only the close association between dogs and humans in the Natufian indeed indicate a special case in human/animal relationship, but that most of the early morphological differences are the same characteristics as found in other domesticated animals in general (Clutton-Brock, 1981; Olsen, 1985; Benecke, 1987; Tchernov & Horwitz, 1991). One of the most interesting phenomena associated with Natufian sites is the complete absence of ‘‘normal’’ wolves. The only known records for large canids have been identified as primitive dogs, characterized by an unproportionally short snout, and hence crowding of teeth, diminution of the carnassials, and a significant decrease of body size (Clutton-Brock, 1981; Olsen, 1985; Benecke, 1987).

Material and Methods Material and comparisons Fossil museum specimens were studied in the palaeontology collection of the Hebrew University of Jerusalem (Hayonim Terrace and Eynan), and in the Natural History Museum, London. It included the material of el-Wad (Bate, 1937), Kebara (TurvillePetre, 1932), and Shukhba (Bate, 1942). The canid material from Neve David (Geometric Kebaran period) (Kaufman & Ronen, 1987) is held by the Department of Prehistory, Haifa University, and the Natufian material from the site of Eynan (=Mallaha) is kept in the Prehistory Museum of Ma’ayan Baruch. Recent museum specimens were based on the canid collection of Tel-Aviv University, Zoological Museum and the zoological collection of the Hebrew University of Jerusalem. The southern Levant wild wolves demonstrate marked pattern of geographic size variation, with a possible gradient from north (Mediterranean ‘‘pallipes’’=Canis lupus pallipes) to south (desert ‘‘pallipes’’=Canis lupus arabs) (Mendelssohn, 1982; Dayan, 1994). The desert wolf was found to be significantly smaller than individuals from the Mediterranean regime of the Levant. Specimens from the Arabian peninsula (but also from southern Israel) may weigh 12 kg, which at present may be considered as the smallest wild wolves. These small sized desert populations were previously used as a reference collection for comparing fossil material of early dogs which originated in northern regions. Some of the characters of the two populations of wolves were found to be non-significant (Dayan, 1994) while others have shown a significant separation. Mendelssohn (1982)

Dogs from the Southern Levant 73

recognized a ‘‘desert’’ pallipes population (occupying regions under 400 mm isohyete) and ‘‘Mediterranean’’ pallipes population (over 400 mm isohyete). However, the separation of ‘‘Mediterranean ‘‘pallipes’’ and desert ‘‘pallipes’’ wolves as different groups seems to be artificial. Due to heavy human occupation and the extensive mortality that the wolf populations suffered during the late historical period in the southern Levant (and elsewhere), a geographical gap between the northern and the southern populations have been created. It is hence assumed that in the recent past the Levantine wolves formed a continuous distribution with a north–south Bergmannian size decrease (Davis, 1981; Davis & Valla, 1978; Dayan, 1994). Obviously, as all the Natufian sites where dogs have been found are located within the Mediterranean regime, they should have been originated from a local, Mediterranean type of wolf. There is now enough evidence to show that the late Pleistocene wolves of the southern Levant were larger than the recent populations (Davis & Valla, 1978; Davis, 1981; Dayan, 1991, 1994). Wolves preceding the Natufian period are extremely rare in the fossil record. In fact the only Geometric Kebaran record of a wild wolf comes from the site in Neve David (Kaufman & Ronen, 1987), while the records of wolves from the Kebaran and Aurignacian of Hayonim cave are doubtful. Careful rechecking of their provenance showed that both records might have originated in Natufian burials (Hayonim cave) which were dug into Kebaran and Aurignacian deposits, hence those records are most probably intrusive. As Dayan (1994) and Dayan et al. (1989, 1990) pointed out, in absence of changes in the structure of the guild (Simberloff & Dayan, 1991) (southern Levantine Canidae in this case), the larger size in one of its members in Natufian, Kebaran or Aurignacian periods may be also an indication for the larger size of others, including wolves of these periods. The Saluki dog was selected for comparison mainly for two reasons: (a) it most probably comprises a very old breed (Przeznziecki, 1984) in the Levant, probably developed either in Mesopotamia or in Egypt, but throughout time it was isolated within this relatively limited geographical region, (b) it comprises a secondary, determinate product of conscious selection for special traits; mainly for high cursoriality and special adaptations for living in arid regions. Comparison with an artificially selected specialized breed, yet well adapted for harsh conditions (and hence partially also naturally selected), facilitates our understanding of the morphological changes which the early Natufian dogs underwent. Unfortunately we found only one ‘‘pure’’ specimen of Saluki in our collections originated in the Sinai desert. The weight of Saluki dogs ranges around 12 kg, and hence is well within the estimated weight of the Natufian dogs, a fact which also facilitates the comparison of the fossil material with this very old, but highly selected, breed.

List of variables Of the three types of landmarks which are usable for epigenetic explanations (Bookstein, 1991: pp. 63–66), we used his ‘‘type 3*’ (external points) only, although the other two landmarks types (Type 1—juxtaposition of tissues; Type 2—maxima of curvature) could have been used for further explanation and better understanding of the biological process which led to the evolvement of dogs from wild wolves. In our case, using the other types of landmarks is useless with such a limited number of individuals. Hence further morphometric approaches will have to wait until more material (of both recent wolves and Natufian dogs) comes to light. Below is a list of cranial, dental and post-cranial measurements. Many of them are standard variables which were intentionally selected to correspond and hence facilitate comparisons with other studies, mainly those of Clutton-Brock et al. (1976), Driesch (1976), and Morey (1992). Unfortunately it was impossible to follow Wayne’s (1986a,b,c) variables for bivariate coefficient analyses as the fossil material never included complete skulls, cranial depth or face length. Maxillary (1) Total length of tooth row (from prosthion to posterior border of last molar alveole in sagittal projection). (2) Total length of tooth row (from prosthion to anterior border of P4 alveole in sagittal. (3) Total length of tooth row (from anterior end of P4 to posterior end of M2 in sagittal projection). (4) Width of incisor row (measured at the alveole border). (5) Width of muzzle (at the canine roots). (6) Length of C1. (7) Width of C1. (8) Length of P2. (9) Width of P2. (10) Length of P4. (11) Width of P4. (12) Length of M1. (13) Width of M1. Mandible (1) Total length of mandible (from posterior border of condyle to distal end of ramus). (2) Total length of tooth row (from posterior border of M3 alveole to distal end of ramus). (3) Total length of mandible minus total length of tooth row. (4) Length of incisor and premolar series (measured at the alveoles). (5) Length of molar series (measured at the alveoles). (6) Depth of ramus behind the canine (measured at the alveoles).

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(7) Depth of ramus (between P/4 and M/1) (measured at the alveoles). (8) Depth of ramus (under middle M/1) (measured at the alveoles). (9) Length of moment arm of temporalis (from back of condyle to top of coronoid process). (10) Length of C1. (11) Width of C1. (12) Length of P2. (13) Width of P2. (14) Length of P3. (15) Width of P3. (16) Length of P4. (17) Width of P4. (18) Length of M1. (19) Width of M1. (20) Length of M2. (21) Width of M2.

Post-cranial skeleton Scapula (1) Greatest length of glenoid region (infra-articular tuberosity to coracoid process). (2) Length of glenoid cavity. (3) Width of glenoid cavity.

(3) Greatest depth of distal end. (4) Greatest width of distal end (across the level of sesamoid fossa).

Metacarpus III (1) Greatest length (base to tip of sagittal crest). (2) Greatest depth of proximal end. (3) Greatest depth of distal end. (4) Greatest width of distal end (across the level of sesamoid fossa).

Metacarpus IV (1) Greatest length (base to tip of sagittal crest). (2) Greatest depth of proximal end. (3) Greatest depth of distal end. (4) Greatest width of distal end (across the level of sesamoid fossa).

Metacarpus V (1) (2) (3) (4)

Greatest length (base to tip of sagittal crest). Greatest depth of proximal end. Greatest depth of distal end. Greatest width of distal end (across the level of sesamoid fossa).

Humerus (1) Greatest length (greater tubercle to capitulum). (2) Greatest length (lesser tubercle to capitulum). (3) Maximal depth of proximal end. (4) Smallest breadth of diaphysis (below nutrient foramen). (5) Greatest width of distal end.

Femur (1) Maximal depth of caput femoris. (2) Greatest width of proximal end (fovea to outer border of greater trochanter). (3) Greater width of distal end (lateral condyle to lateral epicondyle).

Radius

Tibia

(1) Greatest length (styloid process to head of coronoid process). (2) Greatest width of proximal end. (3) Smallest width of diaphysis. (4) Greatest width of distal end.

(1) Greatest length (intercondylar eminence to medial malleolus). (2) Greatest width of proximal end. (3) Smallest width of diaphysis. (4) Greatest width of distal end.

Ulna (1) Smallest anconeal (2) Smallest anconeal

Calcaneus (1) Greatest length. (2) Greatest width.

depth across anconeal notch (between process and olecranon). depth across trochlear notch (between process and coronoid process).

Metatarsus II Metacarpus II (1) Greatest length (base to tip of sagittal crest). (2) Greatest depth of proximal end.

(1) (2) (3) (4)

Greatest length (base to tip of sagittal crest). Greatest depth of proximal end. Greatest depth of distal end. Minimum width of shaft.

Dogs from the Southern Levant 75

Metatarsus V (1) Greatest length (base to tip of sagittal crest). (2) Greatest depth of proximal end. (3) Greatest depth of distal end. (4) Minimum width of shaft. Phalanxx I (1) Greatest length. (2) Greatest width of proximal end. (3) Greatest width of distal end. Phalanx II (1) Greatest length. (2) Greatest width of proximal end. (3) Greatest width of distal end. Phalanx III (1) Greatest length. (2) Greatest width of proximal end. (3) Greatest width of distal end. Phalanx V (1) Greatest length. (2) Greatest width of proximal end. (3) Greatest width of distal end. Statistical procedures We extensively used the bivariate plots displayed as scattergrams in order to visually and individually demonstrate differences between the Natufian population of early dogs with the recent local wolves. As complete skulls from the Natufian do not exist we could not select this measurement as an independent and comparable variable, but used other variables instead, such as total maxillary and mandibular measurements, which are highly correlated to the condylo-basal length of the skull. As Wayne (1986a) stressed a bivariate approach can be more simply and clearly understood in terms of ‘‘size dependent changes in proportions’’ (p. 244). We fully accepted Wayne’s claims that since all cranial measurements are tightly inter-correlated, ‘‘the difference in scaling between dental and cranial measurements’’ (p. 244) will not be appreciably affected by the choice of the independent measurement. Percentiles are values above and below which certain percentages of the variables fall. Percentile plot may be useful in comparing and distinguishing variables of different groups. In order to compare the distribution of our different variables of the recent wolves and the Natufian dogs, we found that using percentile plots allow us to observe the percentage of the data that are less than or equal to an observation. We divided the

total frequency into quantiles. The added reference lines displays the 10th, 25th (quartiles), 50th (quintiles), 75th and 90th percentiles. In this way the percentile ranks of all the numerical values are shown. When the corresponding percentiles of the recent wolves were set against the Natufian dogs, a graphic comparison of the distribution of the two groups were clearly shown. Computing the percentile points of a population enables a good estimation of its distribution curve, when the 50th percentile is the median and the 10th exhibits around 1·25 S.D. of the median; the 25th point—20·75 S.D.; the 75th point—21·25 S.D. and the 90th—21·75 S.D. We found that the best way to test whether two distributions which are independent of each other have been drawn from a unique population, is to use the Kolmogorov–Smirnov two-sample test, as it tests whether the distribution of a continuous variable is the same for two groups; in our case the Natufian dogs against the recent wolves of the southern Levant. It is normally calculated by a comparison of two populations at number of points and than considering the maximum difference between the two distributions. If the two samples belong to a unique population, then the cumulative distributions of both samples (as represented graphically in the percentile plots) may be expected to be fairly close to each other, inasmuch as both samples showed only random deviation from the population distribution. Yet if they are too far apart of any point, it suggests that they belong to different populations. For small samples, as we have in our study, H0 will be rejected if the value of KD (K=the number of scores equal to or less than x, D=maximum vertical deviation—D—of the two cumulative distributions for the largest deviation in the predicted direction), is large enough and the probability associated with the occurrence under H0 is equal to or less than =0·01. As we are comparing two natural distinctive groups of animals (at least by time difference—Natufian dogs versus recent southern Levantine wolves), and because the measurements of one group is essentially controlled by the other, we found it appropriate to use the more powerful paired t-test. The significance level chosen to reject the H0 hypothesis was P=0·01.

Morphological Analysis Mandible Evaluation of the equality/inequality of the sample means of the two groups, the recent wolves with the Natufian dogs, have shown that some morphological traits are indeed different, while others did not change much, or with any statistical significance. A list of all mandibular measurements (range, mean, S.D., standard error, t-test and F-test) is given in Table 1. The most classical characteristic of early domesticated dogs is thought to be the disproportionate shortening of the

76

E. Tchernov and F. F. Valla

Table 1. Range (mm), mean (S.D.), standard error (S.E.), paired t-test comparisons (hypothesized difference=0), F-test (hypothesized difference=1) of mandibular variables of recent wolves and Natufian dogs

Variable

Count

Mean

S.D.

S.E.

(a) Length of C/1

Recent Wolves

13

12·19

0·867 0·241

(b) Length of C/1 (a) Width of C/1

Natufian Dogs Recent Wolves

4 13

11·23 8·16

0·871 0·435 0·598 0·163

(b) Width of C/1

Natufian Dogs

4

7·43

0·837 0·418

(a) Length of P/2

Recent Wolves

13

10·79

1·044 0·289

(b) Length of P/2 (a) Width of P/2

Natufian Dogs Recent Wolves

4 13

9·61 5·18

0·471 0·235 0·362 0·1

(b) Width of P/2

Natufian Dogs

4

4·77

0·104 0·052

(a) Length of P/3

Recent Wolves

12

12·28

0·932 0·269

(b) Length of P/3 (a) Width of P/3

Natufian Dogs Recent Wolves

7 12

11·56 5·70

0·132 0·137 0·354 0·102

(b) Width of P/3

Natufian Dogs

7

5·60

0·271 0·102

(a) Length of P/4

Recent Wolves

13

14·17

1·121 0·311

(b) Length of P/4 (a) Width of P/4

Natufian Dogs Recent Wolves

4 13

12·77 7·01

0·58 0·29 0·624 0·173

(b) Width of P/4

Natufian Dogs

6

6·34

0·49

0·2

(a) Length of M/1

Recent Wolves

13

9·80

0·6

0·166

(b) Length of M/1 (a) Width of M/1

Natufian Dogs Recent Wolves

7 13

8·78 25·22

0·628 0·237 1·388 0·385

(b) Width of M/1

Natufian Dogs

7

22·17

0·873 0·33

(a) Length of M/2

Recent Wolves

13

0·53

0·962 0·267

(b) Length of M/2 (a) Width of M/2

Natufian Dogs Recent Wolves

7 13

9·78 7·56

0·681 0·257 0·721 0·2

(b) Width of M/2

Natufian Dogs

7

7·64

0·718 0·271

(a) Length of Molar Series

Recent Wolves

14

41·04

2·277 0·609

(b) Length of Molar Series

Natufian Dogs

4

37·82

2·655 1·328

(a) Total Length of Tooth Row

Recent Wolves

13

109·70

7·063 1·959

(b) Total Length of Tooth Row

Natufian Dogs

4

99·64

5·734 2·867

(a) Length of Incisor and Premolar Series (b) Length of Incisor and Premolar Series (a) Depth of Ramus between P/4 and M/1 (b) Depth of Ramus between P/4 and M/1 (a) Depth of Ramus under Middle M/1 (b) Depth of Ramus under Middle M/1 (a) Depth of Ramus behind the Canine (b) Depth of Ramus behind the Canine

Recent Wolves

13

74·80

2·952 0·819

Natufian Dogs

4

62·53

3·192 1·596

Recent Wolves

14

26·04

2·15

0·575

Natufian Dogs

4

21·93

1·75

0·875

Recent Wolves

14

26·86

2·322 0·621

Natufian Dogs

2

25·96

0·636 0·45

Recent Wolves

14

18·87

1·978 0·529

Natufian Dogs

2

17·30

0·325 0·23

Mean Diff.

df

t-value

P-value

0·964

15

1·943

0·071

0·991

0·993

0·728

15

1·971

0·068

0·495

0·495

1·179

15

2·155

0·048

4·918

0·127

0·416

15

2·226

0·042

12·142

0·032

0·718

17

1·936

0·069

6·588

0·019

0·097

17

0·621

0·543

1·703

0·475

1·401

15

2·366

0·032

3·731

0·195

0·671

17

2·315

0·033

1·621

0·585

1·022

18

3·577

0·002

0·913

0·901

3·046

18

5·238

<0·0001

2·531

0·212

0·661

18

1·606

0·126

1·996

0·349

18 "0·234

0·817

1·009

0·99

3·217

16

2·412

0·028

0·735

0·749

10·067

15

2·583

0·021

1·517

0·668

12·269

15

7·148

<0·0001

0·855

0·872

4·11

16

3·483

0·003

1·509

0·669

0·897

14

0·529

0·605

13·318

0·054

1·571

14

1·089

0·295

36·996

0·128

"0·079

F-value (V.R.) P-value

Dogs from the Southern Levant 77 46

Length of molar series (mm)

44 42 3a 40

2 1

38 36 3b 34 32 30 55

60

65 70 Length of incisors and premolar series (m)

75

80

Figure 6. Plot of the ratio between the length of incisors and premolar series and the length of the molar series. While the length of the molar series of the Natufian dogs (which actually measures the proximal region of the mandible) greatly overlap the recent wolves, the length of the incisors and premolar series of the dogs (a measures for the distal region of the jaw) is far apart from that of the wolves, clearly showing a conspicuous shortening of the distal part of the snout, but much less in the proximal region, in the Natufian dogs. 1: Hayonim Terrace (dog 1—left); 2: Hayonim Terrace (dog—right); 3a: Shukhba (dog 1); 3b: (Shukhba (dog 2). -: Natufian dogs; /: recent wolves; 4: Saluki dog.

Obviously when the total length of the tooth row is compared between dogs and wolves. Differences are less visible when compared with the most affected part of the mandible (Table 1), or the tip of the snout. The Saluki dog, however, does not show such a degree of 85

80

75 Value

muzzle, which is expressed in marked reduction of the maxillary and the mandibular length (Clutton-Brock, 1962; Olson, 1985; Benecke, 1987). But if we divide the snout, or the maxillary and the mandible into anterior (incisors+premolars series) and posterior (molar series) portions, the results show marked differences in the rate and amount of changes between the two regions of the snout. The main changes in the length of the snout, as is known in other groups of vertebrates (Tchernov, 1986) was mainly expressed in the anterior part of the snout, while the posterior region undergoes as a rule only slight changes. This trend of change is clearly evidenced in Figure 6, in the percentile plot (Figure 7) and in Tables 1 & 2, which clearly indicate a conspicuously shortening of the anterior part of the mandible (along the incisors and the premolar teeth), in comparison with the posterior portion of the lower jaw (along the molar series). One of the Shukhba specimens (Figure 6, specimen 3b) shows the most extreme shortening of the snout, indicating a progressive selective relaxation in this trait as compared with wild wolves. As it is shown in Table 1, the shortening of the anterior part of the ramus along the incisors and the premolar teeth, is the most affected trait. If the depth of the ramus (measured under C1) is taken against the total length of the mandible (Figure 8) the dog of Hayonim Terrace (the only specimen from which both measurements exist) shows a mark shortening of the lower jaw. Yet this size reduction, as seen above, is mainly caused by the shortening of the distal region, moderately in its middle part, and only slightly in the proximal region of the mandibular ramus.

70

65

60

55

0

20

40 60 Percentile

80

100

Figure 7. A percentile plot of the length of incisors and premolar series in Natufian dogs and recent wolves from the southern Levant. The dogs and the wolves show very different values, indicating the drastic shortening of the anterior part of the snout in the Natufian dogs. ,: Length of incisor and premolar series (recent wolf ); .: length of incisor and premolar series (Natufian dog).

Length mand. (dog) * * 138·750 * *

Length tooth row (dog) * 95·360 101·165 103·910 *

Length molar series (dog) * 35·790 38·475 39·845 *

L. inc.+permol. ser. (dog) * 60·025 61·995 65·040 * L. md.–L. tooth row (dog) * * 39·210 * * L. moment arm temp. (dog) * * 38·670 * *

Length mand. (wolf) 156·819 161·240 162·860 172·090 174·893

Length tooth row (wolf) 98·368 104·123 110·340 116·264 117·406

Length molar series (wolf) 38·332 39·890 40·985 41·540 44·039

L. inc.+premol. ser. (wolf) 71·530 72·707 74·030 78·065 78·380

L. md.–L. tooth row (wolf) 46·224 49·480 53·980 61·630 68·055

L. moment arm temp. (wolf) 41·194 44·860 48·145 53·250 55·339

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

Width of P/3 (wolf) 5·349 5·495 5·645 5·760 6·273

Length of P/3 (wolf) 10·992 11·710 12·600 12·890 13·293

Width of P/2 (wolf) 4·704 4·787 5·060 5·355 5·576

Length of P/2 (wolf) 9·450 10·283 10·840 11·385 11·830

Width of C/1 (wolf) 7·222 7·747 8·37 8·61 8·756

Length of C/1 (wolf) 11·07 11·405 12·37 12·815 13·456

Width of P/2 (dog) 5·246 5·365 5·700 5·840 5·864

Length of P/3 (dog) 10·969 11·410 11·495 11·680 11·833

Width of P/2 (dog) * 4·710 4·720 4·820 *

Length of P/2 (dog) * 9·240 9·550 9·980 *

Width of C/1 (dog) * 6·85 7·495 8·005 *

Length of C/1 (dog) * 10·665 11·535 11·79 *

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

Depth mand. (behind C/1) (wolf) 16·580 17·440 18·675 20·590 21·320 Depth mand. (P/4 & M/1) (wolf) 23·095 24·480 26·000 28·220 28·521

Depth mand. (under M/1) (wolf) 23·346 25·030 27·310 28·550 29·711

Width of M/1 (wolf) 22·972 27·770 25·400 26·107 26·666

Length of M/1 (wolf) 9·146 9·467 9·780 10·025 10·702

Width of P/4 (wolf) 6·280 6·625 6·900 7·250 7·920

Length of P/4 (wolf) 12·474 13·238 14·59 14·795 15·274

Depth mand. (behind C/1) (dog) * 17·070 17·300 17·530 * Depth mand. (P/4 & M/1 (dog) * 20·490 22·040 23·360 *

Depth mand. (under M/1) (dog) * 25·510 25·960 26·410 *

Width of M/1 (dog) 20·912 21·490 22·240 22·833 23·184

Length of M/1 (dog) 7·880 8·275 8·920 9·185 9·518

Width of P/4 (dog) 5·500 6·400 6·450 6·600 6·780

Length of P/4 (dog) * * 13·0212·42 13·12 *

Table 2. Percentile table reporting the 10th, 25th, 50th, 75th and 90th percentile points of different variables of Natufian dogs and recent wolf populations

Width of M/1 (wolf) * * 28·820 * *

Length of M/1 (wolf) * * 11·000 * *

78 E. Tchernov and F. F. Valla

Dogs from the Southern Levant 79 180

Total length of mandible (mm)

175 170 165 160 155 150 145

1 2

140 135 130 14

15

16

18 19 17 Depth of ramus behind canine (mm)

20

21

22

Figure 8. Plot of the ratio between the depth of the ramus behind canines and the total length of mandible. While the depth of the ramus remains unchanged in the Natufian dogs, this plot shows the conspicuous shortening of the mandible. 1: Hayonim Terrace (dog 1—left); 2: Hayonim Terrace (dog 1—right). /: Natufian dogs; -: recent wolves; 4: Saluki dog.

snout shortening, but as with many other traits, this trend could have been secondarily reversed much later under directional selection. The depth of the mandibular ramus behind the canine, as seen in Figure 8 was practically unchanged. This is probably due to the fact that the size of the canines were only little affected. Therefore in order to firmly hold and support the relatively larger canine teeth (Figure 8, specimen 3) the depth of the ramus in this region cannot be changed. The critical shortening of the anterior portion of the mandible is clearly followed by a marked and significant decrease in the length of P3 (Tables 1 & 2; Figure 9) in relation with the little affected depth of the ramus, especially the region under M1. The Saluki dog and the Eynan (=Mallaha) specimen show a most advance reduction of this tooth. The extreme reduction in the size of P3 in the Saluki dog (as an old breed) suggests a further differentiation of this tooth in later periods. The Eynan specimen is the only Natufian dog which shows a marked thinning in the depth of the ramus under M1, while all the other Natufian dogs, including the Saluki dog, do not display any change in the thickness of the ramus in this region. The depth of the mandible was differently changed at different sections along the ramus. The most severe decrease in the depth of the ramus took place under P4- M1,and posteriorly (Tables 1,2 & 3 and Figure 10), while almost no change could have been detected in the depth of the ramus either under the canines or under M1. In term of thickness the anterior part of the mandible seems to be less affected than its posterior portion (behind M1). Therefore, the Natufian dogs show a maximum shortening in the anterior part of the

snout, but little change in the depth of the ramus in this region. As shown above, however, the posterior part of the ramus, however, did not undergo significant shortening, yet the depth of the ramus at this region became significantly slender. Functionally it indicates a less effort for mastication (relying on softer food) in the Natufian dogs, compared with recent wolves. The most ‘‘delicate’’ jaws are found in one of the three known specimens from the site of Shukhba. The relatively long muzzle (but relatively low ramus) of the Saluki dog can again be explained as a reversed secondary selection. The relative constancy of the canine size (and shape) of the domesticated wolves (and other canids) may not necessary due to functionality, but stem from some developmental constraints. This organ emerges relatively early in the ontogeny of the canids, and hence any morphological change in the canines may interfere with too many other morphological characters which might be deleterious if also changed. As the least affected tooth is C1 (Tables 1, 2 & 3), which shows no notable reduction in size and proportion, its root should have been also remained unchanged in order to maintain the same size and shape of the crown. Indeed the depth of the ramus at this region has been left unchanged. P2 and P3 were only slightly reduced, while M1 underwent a severe decrease in size in spite of the fact that this part of the ramus became only slightly shorter (Table 1,2 & 3). A comparison with the length, but particularly with the width of M1 with the relatively unchanged C1, clearly show a conspicuous decrease in the size of this tooth (Figure 9). If the area of M1 is calculated (as shown in legend of Figure 9) those differences between the dogs and the wolves become even more conspicuous. The

80

E. Tchernov and F. F. Valla 30 I

29 28 Width of M/1 (mm)

27

a c

26

b

25 24

2

23

3

22 21 20 7.5

1b

2

4

1a 4

8

8.5

9

9.5 10 Length of M/1 (mm)

10.5

11

11.5

Figure 9. Plot of the ratio between the Length of M1 and Width of M1. The approximation values of the occlusion area of M1 of the dogs and some of the wolves is given as well, showing a better correlation between the size of the animal and the size of the molar. There is a proportional decrease in the size of M1 and the weight of the animal. 1: Hayonim Terrace (dog 1); 2: Hayonim Terrace (dog 2); 3: Eynan; 4: Shukhba. I: (L#W)=316·8; a: (L#W)=275·6; b: (L#W)=242·25; c: (L#W)=253·50; 1a: (L#W)=213·29; 1b: (L#W)=219·65; 2: (L#W)=173·97; 2: (L#W)=196·24; 3: (L#W)=191·71; 4: (L#W)=162·53. a: 24·2 kg; b: 28 kg; c: 23·2 kg. /: Natufian dogs; -: recent wolves; 4: Saluki dog; ;: Geometric Kebaran wolf (Neve David).

13 12.5

Length of P/2 (mm)

12

Golan Height 25 kg

11.5 11 10.5

1

10

1 3

9.5

2

9 8.5 8 4.2

4.4

4.6

4.8

5 5.2 Width of P/2 (mm)

5.4

5.6

5.8

Figure 10. Plot of the ratio between the width of P2 and length of P2. There is an obvious diminution in the size of the tooth in the Natufian dogs, but in particular in the Saluki dog. 1: Hayonim Terrace (dog 1); 2: Eynan; 3: Shukhba. /: Natufian dogs; -: recent wolves; 4: Saluki dogs.

consequence of this differential morphometric change in M1 cannot lead to any crowding of the molars, but vice versa, leaving even more space between the teeth. If any crowding of the teeth would have taken place it would have shown within the region of the premolars and the canines, where the teeth are relatively much

less shortened, but the ramus along this region was markedly reduced. On the other hand the molar series were reduced in size, yet that part of the ramus did not undergo marked changes, hence there was no crowding of the teeth also in the rear part of the ramus. The most affected tooth of the Natufian dogs is therefore M1,

Dogs from the Southern Levant 81 Table 3. Kolmogorov–Smirnov test for all the mandibular, maxillary and post cranial variables of southern Levantine recent wolves and Natufian dogs Variables

df

Dog (n)

Wolf (n)

Max. Diff.

Chi Square

P-value

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2

4 4 4 4 7 7 4 6 7 7 1 4 4 4 1 2 2 4 1

13 13 13 13 12 12 13 13 13 13 14 13 14 13 14 14 14 14 14

0·615 0·519 0·769 0·769 0·750 0·321 0·769 0·692 0·780 0·846 1·000 0·769 0·714 1·000 1·000 0·571 0·714 0·857 1·000

4·633 3·299 7·240 7·240 0·000 1·827 7·240 8·870 11·079 13·031 3·733 7·240 6·349 12·235 3·733 2·286 3·571 9·143 3·733

0·1972 0·3844 0·0536 0·0536 0·0222 0·8022 0·0536 0·0391 0·0079 0·0030 0·3093 0·0536 0·0836 0·0044 0·3093 0·6378 0·3354 0·0207 0·3093

2 2 2 2 2 2 2 2 2 2 2 2 2

3 7 2 3 6 4 8 4 1 1 3 3 4

11 12 11 13 13 13 13 13 13 13 12 11 13

0·727 0·274 0·727 0·846 0·333 0·750 0·500 0·846 1·000 0·923 0·250 0·667 0·346

4·987 1·326 3·580 6·981 1·825 6·882 4·952 8·760 3·714 3·165 0·600 4·190 1·466

0·1652 <0·9999 0·3339 0·0610 0·8032 0·0641 0·1681 0·0250 0·3122 0·4110 <0·9999 0·2461 0·9609

2 2

3 3

6 6

0·833 1·000

5·556 8·000

0·1244 0·0366

2 2

2 2

4 4

1·000 1·000

5·333 5·333

0·1390 0·1390

2 2 2 2 2

3 2 1 2 1

6 6 6 6 6

0·833 0·833 1·000 1·000 1·000

5·556 4·167 3·429 6·000 3·429

0·1244 0·2490 0·3602 0·0996 0·3602

2 2 2 2

2 1 2 1

4 4 4 4

1·000 0·750 1·000 1·000

5·333 1·800 5·333 3·200

0·1390 0·8131 0·1390 0·4038

2 2

3 1

4 4

1·000 1·000

6·857 3·200

0·0649 0·4038

Mandible Length of C1 Width of C1 Length of P2 Width of P2 Length of P3 Width of P3 Length of P4 Width of P4 Length of M1 Width of M1 Total length of mandible Total length of tooth row Length of molar series Length of incisors and premolar series Total length of mandible minus total length of tooth row Depth of ramus under M1 Depth of ramus behind canines Depth of ramus between P4 and M1 Length of moment arm of temporalis Maxillary Length of C1 Width of C1 Length of P2 Length of P4 Width of P4 Length of M1 Width of M1 Total length of tooth row (prosthion to last alveole) Total length of tooth row (prosthion to anterior border of P4) Length of tooth row Width of incisor row Width of muzzle (at canine roots) Minimum width of muzzle (between P1 and P2) Femur Maximum depth of caput femoris Greatest width of proximal end Scapula Greatest length of glenoid region Length of glenoid cavity Humerus Greatest width of distal end Smallest breadth of diaphysis Maximal length of proximal end Greatest length (lesser tubercle to capitulum) Greatest length (greater tubercle to capitulum) Radius Greatest width of distal end Smallest breadth of diaphysis Maximal length of proximal end Greatest length Ulna Smallest depth across trochanter notch Smallest depth across anoconeal notch

and in its width more than its length. The net effect is hence an overall narrowing of the lower molars. The same change is shown also in the Saluki dog. Worth noting is the very wide range of variability in M1 within the Natufian population of dogs, particularly shown by the Shukhba specimens (Figure 9). M2 shows very little changes in spite of the notable decrease in body size, and the shortening of the snout (Tables 1, 2 & 3). The range of variability, however, of the Natufian dogs is extremely wide, particularly shown by one of the Shukhba specimens. Worth mentioning is the small size of the M2 of the Saluki dog.

P2 shows a proportional decrease in both the length and width of the tooth, in comparison with the recent population of wolves (Figure 10, Tables 1, 2 & 3). This may reflect the significant shortening of the anterior part of the mandibular ramus. This fact also shows that there is no ‘‘crowding’’ of the teeth, at least in these early Natufian dogs, but a rather simultaneous reduction of the premolars (excluding the canines) and the distal region of the jaws. The Saluki dog shows the smallest P2, which is probably the result of long selective relaxation in the shape and the size of the molars, with no connection, or dependence, to its

E. Tchernov and F. F. Valla

Length of tooth row (from anterior end of P\4 to posterior end of M/2 in sagittal projection) (mm)

82

43 42 41 40

1

39 2

38 37 36

3

35 34 33 95

100

105

115

110

120

125

Total length of tooth row (from prosthion to posterior border of last molar alveole in sagittal projection) (mm)

Minimum width of muzzle (between P\1 and P\2) (mm)

Figure 11. This plot displays the ratio of the total length of tooth row against its posterior portion (from anterior end of P4 to posterior end of M2), showing that the posterior region underwent only little changes in its length (compared with the distal portion of the maxillary). 1: Hayonim Terrace (dog 1); 2: Kebara; 3: el Wad. -: Natufian dogs; / recent wolves; 4: Saluki dog.

48 46

C. lupus – Golan Heights 29.35 kg

44 2 4

42 40 38

1

3 36 34 32 95

100

105

110

115

120

125

Total length of tooth row (mm) Figure 12. Plot of the ratio between the total length of tooth row and the minimum width of muzzle (between P1 and P4), showing the constant breadth of the snout in dogs and wolves, while its length was radically shortened. 1: Hayonim Terrace (dog 1); 2: Hayonim Terrace (dog 2); 3: Kebara; 4: el Wad. /: Natufian dogs; -: recent wolves; 4: Saluki dog.

secondary lengthening of the snout due to reversal selection. There is also quite a notable decrease in the size of P4 in the Natufian dogs, which displays a huge variability, as shown by the three specimens from the site of Shukhba; one of them possessed the smallest

P4, while the two others were only slightly affected. The Saluki dog is well within the size of the population of Natufian dogs (Tables 1, 2 & 3). The three represented dogs display an immense range of variability. The shortest snout is found in the dog of Kebara.

Dogs from the Southern Levant 83

Length of tooth row (from prosthion to anterior border of P\4 alveole) (mm)

85 Golan Height 29 kg 80

75 1 3

70

2 65

60 95

100

105

115

110

120

125

Total length of tooth row (from prosthion to posterior border of molar alveole) (mm)

Minimum width of muzzle (between P\1 and P\2) (mm)

Figure 13. Plot of the ratio of total length of tooth row between prosthion to posterior border of molar alveole and prosthion to anterior border of P4 alveole, showing the allometric relationship between the total length of the jaws to its anterior portion. 1: Hayonim Terrace (dog 1); 2: Kebara; 3: el Wad; /: Natufian dogs; -: recent wolves; 4: Saluki dog.

50 48 46 C. lupus – southern Negev ≈ 14 kg

44 1

3

42 40

1

38 2 36 34 32 30 19

19.5

20

20.5

21

21.5

22

22.5

23

23.5

24

Length of P\4 (mm) Figure 14. This plot displays the ratio between the length of P4 and the width of muzzle (between P1 and P2). While the width of the snout remains unchanged in the Natufian dogs, the length of P4 markedly decrease in size synchronously with the shortening of the snout (see Figure 19). 1: Hayonim Terrace (dog 1); 2: Kebara; 3: el Wad. /: Natufian dogs; -: recent wolves; 4: Saluki dog.

Maxillary The width of the snout did not actually change in the Natufian dogs. This is clearly shown by the measurements of the width of the muzzle taken along different sections of the snout. At the section of the canines and between P1 and P2 the width appears to remain fairly constant (Figure 11, Tables 4, 5 & 6). The general

shortening of the snout, measured as the total length of the tooth row (from prosthion to posterior border of last molar) against the width of the muzzle (at the canine roots), is notable in Figure 15. As expected, there is a conspicuous decrease in the length of the anterior portion of the snout, while its posterior region was only slightly changed as exemplified in Figures 13 & 14. In fact both the width of the snout and its

Total length tooth row (pros. P/4) (dog) * 100·475 104·480 107·010 * Length tooth row (dog) * * 64·880 * * Width muzzle (at canine roots) (dog) * 38·583 42·910 42·910 * Min. width muzzle (P/1 & P/2 (dog) * 36·660 39·480 41·835 * Width of incisor row (dog) * 26·898 27·760 28·375 *

Total length tooth row (pros.-P/4) (wolf) 107·366 111·118 114·080 117·375 118·976

Length tooth row (wolf) 72·140 72·435 74·430 77·100 80·062

Width muzzle (at canine roots) (wolf) 37·420 38·075 40·690 40·913 42·086

Min. width muzzle (P/1 & P/2) (wolf) 36·860 38·662 40·790 42·207 44·132

Width of incisor row (wolf) 29·075 26·820 27·500 28·820 29·351

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

Width of P/2 (wolf) 4·410 4·908 5·220 5·410 5·454

Length of P/2 (wolf) 10·802 12·212 12·460 13·140 13·524

Width of C/1 (wolf) 6·679 6·970 7·445 7·950 8·112

Length of C/1 (wolf) 10·932 11·438 12·120 12·663 13·012

Width of P/2 (dog) * 4·480 4·700 4·920 *

Length of P/2 (dog) * 7·470 7·610 7·750 *

Width of C/1 (dog) 6·770 7·028 7·250 7·745 7·790

Length of C/1 (dog) * 11·012 11·020 11·462 *

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

10 25 50 75 90

Width of M/1 (wolf) 17·180 17·513 18·460 19·143 20·044

Length of M/1 (wolf) 14·158 14·443 15·030 15·620 15·932

Width of P/4 (wolf) 11·606 11·877 11·990 12·480 13·192

Length of P/4 (wolf) 20·770 22·250 22·840 23·153 23·524

Table 4. Percentile table reporting the 10th, 25th, 50th, 75th and 90th percentile points of different variables of Natufian dogs and recent wolf populations

Width of M/1 (dog) 15·435 16·265 17·340 19·025 20·555

Length M/1 (dog) * 13·095 13·280 14·490 *

Width of P/4 (dog) 10·340 11·420 12·115 12·480 12·714

Length of P/4 (dog) * 20·250 20·490 21·067 *

84 E. Tchernov and F. F. Valla

Dogs from the Southern Levant 85 Table 5. Range (mm), S.D., standard error (S.E.), paired t-test comparisons (hypothesized difference=0, F-test (hypothesized difference=1), of maxillar variables of recent wolves and Natufian dogs

Variable

Count

Mean

S.D.

S.E.

(a) Length of C/1

Recent Wolves

11

12·050

0·779 0·235

(b) Length of C/1 (a) Width of C/1

Natufian Dogs Recent Wolves

3 12

11·213 7·443

0·344 0·198 0·57 0·160

(b) Width of C/1

Natufian Dogs

7

7·347

0·46

0·170

(a) Length of P/2

Recent Wolves

11

12·420

1·26

0·380

(b) Length of P/2 (a) Width of P/2

Natufian Dogs Recent Wolves

2 11

7·610 5·063

0·2 0·140 0·484 0·150

(b) Width of P/2

Natufian Dogs

2

4·700

0·311 0·220

(a) Length of P/4

Recent Wolves

13

22·533

1·027 0·285

(b) Length of P/4 (a) Width of P/4

Natufian Dogs Recent Wolves

3 13

20·640 12·309

0·56 0·323 0·786 0·218

(b) Width of P/4

Natufian Dogs

6

11·848

0·915 0·373

(a) Length of M/1

Recent Wolves

13

15·031

0·642 0·178

(b) Length of M/1 (a) Width of M/1

Natufian Dogs Recent Wolves

4 13

13·793 18·415

1·164 0·582 1·02 0·282

(b) Width of M/1

Natufian Dogs

8

17·630

1·953 0·691

(a) Total length of tooth row (prosthion to posterior border of M/3) (b) Total length of tooth row (prosthion to posterior border of M/3) (a) Length of tooth row (ant. border of P/4 to post. border of M/2) (b) Length of tooth row (ant. border of P/4 to post. border of M/2) (a) Length of tooth row (prosthion to anterior border of P/4) (b) Length of tooth row (prosthion to anterior border of P/4)

Recent Wolves

13

114·107

4·393 1·218

Natufian Dogs

4

103·742

4·503 2·252

Recent Wolves

13

39·550

1·890 0·520

Natufian Dogs

1

35·630

0

Recent Wolves

12

114·690

1·788 0·539

Natufian Dogs

3

103·070

3·331 1·923

(a) Width of incisor row (measured at the alveole border) (b) Width of incisor row (measured at the alevole border) (a) Width of muzzle (at the canine roots) (b) Width of muzzle (at the canine roots) (a) Minimum width of muzzle (between P/1 and P/2) (b) Minimum width of muzzle (between P/1 and P/2)

Recent Wolves

12

27·721

1·225 0·354

Natufian Dogs

3

27·650

0·989 0·571

Natufian Dogs

11

39·814

1·788 0·539

Natufian Dogs

3

40·987

3·331 1·923

Recent Wolves

13

40·635

3·044 0·844

Natufian Dogs

4

39·248

3·066 1·533

posterior portion remain unchanged in all Natufian dogs. This is not seen in the Saluki dog, and may be explained as a secondary reversed selection for a longer snout. The width of the incisor row is the least affected variable in these dogs. The P-value of the t-test (Table 3) and the percentiles (Table 4), and its maximum difference tested by Kolmogorov–Smirnov test (Table 3) show that there is actually no difference between the two sampled populations. Figure 14 clearly exemplifies the stability of the width of the muzzle, in this case,

Mean Diff.

df

t-value

F-value P-value (V.R.) P-value

0·430

2

3·279

0·082

3·15

0·100

0·010

6

0·032

0·975

0·146

0·707

0·450

1

17·481

0·036

40·506

0·122

0·380

1

9·500

0·068

2·421

0·465

2·077

2

3·207

0·085

3·359

0·252

0·167

5

0·387

0·715

1·353

0·308

0·782

3

3·836

0·032

7·806

4·543

0·782

7

0·868

0·414

3·694

0·023

7·683

3

3·718

0·034

—

—

—

—

—

—

—

—

—

—

—

—

16·85

0·000

0

"0·890

2 "4·421

0·048

1·533

0·460

"3·087

2 "1·897

0·198

0·718

0·413

"1·115

3 "4·421

0·048

1·014

0·420

across the minimum width, with no relation with the size of the animal. The most elongated maxillaries are hence found in the largest wolves. The Natufian dogs and the recent wolves are very similar in the width of the snout which agrees well with Wayne’s (1986a,c) argument, that the width of the snout in dogs did not change, and therefore became relatively wider in infants, or in individuals with a smaller body size. The Natufian dogs, while undergoing significant size reduction of the distal region of the snout, is a typical paedomorphic phenomenon.

86

E. Tchernov and F. F. Valla

Table 6. Assorted t-test values of 12 maxillary variables arranged from the most affected variables to the least ones Variable 1 2 3 4 5 6 7 8 9 10 11 12

Width of incisor row (measured at the alveole borders) Width of muzzle (at the canine roots) Minimum width of muzzle (between P/1 and P/2) Width of C/1 Width of P/4 Width of M/1 Length of P/4 Length of C/1 Total length of tooth row (prosthion to last molar alveole) Length of M/1 Width of P/2 Length of P/2

t-test

P-value

"4·421

0·9752

"1·897 "0·2

0·715 0·5793

0·032 0·387 0·868 3·207 3·279 3·718

0·414 0·1983 0·085 0·0818 0·0668 0·0476

3·836 9·5 17·481

0·0364 0·0339 0·0312

13.5 13.2 13 12.8

Value

12.5 12.2 12 11.8 11.5 11.2 11 10.8

0

20

40 60 Percentile

80

100

Figure 15. A percentile plot of the length of C1 in Natufian dogs and recent wolves, showing much the same values in both forms, indicating that the canines remained practically unchanged in the Natufian dogs in spite of the great reduction of the anterior portion of the snout. ,: Length of C1; .: length of C1.

Together with the severe shortening of the snout compared with the relatively stabile width of the snout (Tables 3, 4 & 5, Figures 16 & 17), especially its anterior portion, P2 and M1 underwent a reduction in size, and here again no crowding of the teeth can be claimed. Figure 11 shows the proportional and the high correlation of the snout length and the size of the animal; the smaller the animal the shorter the snout. The Natufian dogs and the smallest recent wild wolves show the same short proportion of the muzzle. The general shortening of the snout in the Natufian dogs, as shown in Figure 12, suggest that this change was more or less proportional to the decrease in the size of both the length and the width of the carnassials. The length

of the snout and the size of the carnassials are, hence, proportional to the size of the animal. In this case the dog from Kebara displays the shortest maxillary, while the decrease of the carnassials were lagging behind. Thus while the width of the muzzle remains unchanged in the Natufian dogs, it is the length of the anterior portion of the snout which is markedly affected. Naturally, the same amount and proportion of shortening along the upper and lower jaws must have taken place for the sake of occlusion, although (see Conclusions) within a certain limit a differential size decrease of the molars is permitted without disturbing the functional occlusion of the teeth. While the width of the muzzle remains unchanged all along in the Natufian dogs, the size of the carnassial greatly decrease in size, proportionally with the changes in body size. This is in particularly shown in one individual of a wild wolf from the arid region of Israel (Figure 14) which weighs 14 kg, and displays the same size of carnassials as a Natufian dog. Hence there is no sign of crowding of the teeth in the anterior part of the snout. The width of incisor row also remains almost unchanged in relation to the shortening of the snout, which indicates once more that the width of the snout remains unaffected (Tables 3, 4 & 5). This is also shown when measuring the relative width of the snout across the canine roots. The width of the muzzle remains unchanged all along (Tables 3, 4 & 5). The conspicuous reduction in the size of the carnassials of the Natufian dogs can be explained by the great deal of relaxation in the selection pressure for efficient predation which, if so, must be connected to at least some amount of dependence on humans. In this case it is more difficult to explain the relative constancy of the canine size. Figure 14 also clearly shows the uniqueness of the Natufian dogs in comparison with the recent wolves. Bearing in mind that the Natufian wolves (like the Geometric Kebaran ones (Dayan, 1994) were much larger than the present southern Levantine populations, the decrease in the body size of the Natufian dogs should have been indeed dramatic. In the lower jaw the canines were only slightly reduced compared with M1 and P2. In particular the width of the canines remained almost unchanged in relation to body size, or the rostral length (Figure 15). Similarly the upper canines remained unchanged in the dogs hence in comparison with P2 and M1, they seem to be disproportional larger. The constancy of the canines size enables a comparison of other changeable elements. Obviously P4 was significantly reduced in size compared with C1, and again clearly places the Natufian dogs—as shown in Figure 16—as a unique group. The Saluki dog shows a reduction both in P4 and C1. Similarly there is also a significant decrease, both in length and width of P2 (Figure 17) and stays in a high correlation with the body size. Note that a 12 kg wolf from the arid region of Israel reaches almost the same premolar size as the Natufian dogs.

Dogs from the Southern Levant 87 24 23.5

Length of P\4 (mm)

23 22.5 22 4

21.5 21

2 4

20.5

3

20 1 19.5 19 10

10.5

11

11.5

12 12.5 Length of C\1 (mm)

13

13.5

14

Figure 16. When the constant C1 is plotted against P4 it shows the conspicuous reduction of the last premolar in all Natufian dogs, as well as in the Saluki dog. 1: Hayonim Terrace (dog 1); 2: Hayonim Terrace (dog 2); 3: Kebara; 4: el. Wad. /: Natufian dogs; -: recent wolves; 4: Saluki dog.

6

Width of P\2 (mm)

5.5

1

5

2

4.5

C. lupus – Dead Sea (12. kg) 4

3.5

7

8

9

10

11 12 Length of P\2 (mm)

13

14

15

Figure 17. Plot of the ratio of width and length of P2. The two Natufian dogs where P2 was preserved display a distinctive decrease in the size of the premolars compared with wolves. Note, however, that a 12 kg recent wolf approaches the size of P2 of a Natufian dog. 1: Hayonim Terrace (dog 1); 2: Kebara. -: Natufian dogs; /: recent wolves.

Post-cranial elements Up until now we know of only two specimens from the Natufian period where post-cranial elements were exposed, identified and measured. Although this is the first time we looked at limb bones of Natufian dogs, statistically it is difficult to signify quantitatively the amount of changes which these dogs underwent in

their post-cranial elements. Yet, even if statistically untested, some of them do show clear cut changes in comparison with recent wolves. Humerus There seems to be a proportional decrease in the size of the humerus with body size as is reflected in the only

88

E. Tchernov and F. F. Valla

Figure 18. Plot of the ratio between the smallest breadth of diaphysis and the greatest width of distal end of the humerus. The marked reduction of the diaphysis and the width of the distal articulation of the humerus seems to be the result of reduced locomotion in the Natufian dogs. 1: Hayonim Terrace (dog 2); 2: Hayonim Terrace (dog 1). /: Natufian dog; -: recent wolves.

Radius—greatest width of proximal end (mm)

22.00

21.00

20.00

19.00

18.00

17.00

16.00 20.00

2

1

21.00

24.00 25.00 22.00 23.00 Radius—greatest width of distal end (mm)

26.00

27.00

Figure 19. This plot displays the ratio between the greatest width of distal end and the greatest width of proximal end of the radius, showing a drastic reduction of the articular facets which may due to reduced mobility in the Natufian dogs. 1: Hayonim Terrace (dog 1—right); 2: Hayonim Terrace (dog—left). /: Natufian dog; -: recent wolves; 4: Saluki dog.

known specimen from Hayonim Terrace. Notice that the very small wolf from the Sinai desert (12·3 kg) approaches the size of a Natufian dog (Figure 18). The maximal depth of the proximal end, however, was only slightly reduced in proportion with the reduction in body size, while the greater width of the distal end was disproportional reduced compared with the width of

the shaft (Tables 3, 7 & 8). The relatively narrow trochlea and capitulum (in dogs from Hayonim Terrace), which form the humeral condyle, indicate a lesser attachment area for the flexors and extensors of the carpus and its digits, probably due to reduced mobility. The two dogs from Hayonim Terrace show especially thin diaphysis in humeri which is different

Dogs from the Southern Levant 89

the proximal and the distal ends in the Natufian dogs, leaving them far apart even from the smallest known recent wolves (Figure 19, Tables 7 & 8). The decrease in size of the radius went much further than the decrease in body size and the proportional diminution of the humerus. Yet, here again, the Saluki dog shows an obvious disproportional enlargement of the radius probably due to its artificial selection for cursoriality.

2.0

Length P/4 (cm)

1.8

1.6 1.4

Y = 9.644 + 0.207

1.2

1.0

0.8

1.3

1.8 2.3 Weight (kg)

2.8

3.3

Figure 20. Correlation between P4 and body weight in recent Levantine wolves, within extrapolation of the weight of the Natufian dogs, estimated to be around 12 kg, hence generally smaller even from the smallest recent wolves from the southern desert of the Levant. Count (Natufian dogs)=4; count (recent wolves)=6; r=0·948; R#2=0·899; standard error=0·849; t-value=11·345; P-value=0·0003. ,: Recent southern Levantine wolves; -: Natufian dogs.

from all recent wolves; even the smallest one which weighs 212 kg, much like the Natufian dogs (see next chapter). Scapula There is a proportional decrease in the size of the scapula in the Natufian dogs (Figure 26). The reduction in size of the glenoid region is proportional to the reduction of its body size. The Saluki dog, however, having very long feet, shows, in relation to the wolves, a disproportional enlargement of the scapula. The greater length of the glenoid region, although smaller, were proportionally reduced in the Natufian dogs. The glenoid cavity itself (its length and its width) also show proportional reduction in size. Ulna There is a disproportional reduction of the anconeal region in the Natufian dogs. The Saluki dog, however, having relatively very long feet, shows, compared with the wolves, a disproportional enlargement of the olecranon region, and in a high correlation with the size of the humerus, but the restricted number of specimens makes it impossible for any effective statistical test (Tables 7 & 8). The disproportional reduction in this region indicates a reduction in the area of attachment for some of the muscles, such as the flexor carpi ulnaris, the anconeus, and part of the triceps, suggesting a reduced functionality. Radius No complete radii were found, but the epiphyseal regions show significant reduction in the width of both

Femur There is a proportional decrease in the size of the Natufian dog from Hayonim Terrace. Maximal depth of caput femoris does not show significance differences, and was reduced in size proportionally to the decrease in body size. But the greater width of the proximal end was disproportional reduced (Tables 3, 7 & 8) in comparison with the caput femoris; or the thickness of the femoral shaft (which was only slightly reduced in diameter), or the insignificant reduction (Tables 3, 7 & 8) of the greatest width of the distal end of the femur. The femora of the only two existing Natufian dogs with post cranial elements clearly indicate that the distance from the fovea (=fovea capitis femoris) to the greater trochanter (=trochanter major) is very narrow. Functionally it means that the attachment area of the middle gluteal muscle (musculus gluteus medius) is relatively much smaller than in wolves, apparently indicating restricted locomotive capabilities, or reduction in mobility, in early dogs. Tibia The decrease in size of the tibia is in full proportion with the changes of the femur (Table 3, 7 & 8). While the Natufian dogs show no difference from the wild wolves, the effect of intentional selection is well shown in the Saluki dog which shows a conspicuous disproportional enlargement of the tibia, a clear adaptation for cursoriality. The distal end of the tibia shows a proportional diminution in size, however small number of measured specimens do not permit any satisfactory statistical tests. Metapodials It is difficult to come to any conclusion about the tarsalia and carpalia of the Natufian dogs, as there are too few remains, as well as too scarce recent comparative material. The only thing we can say is that all the measurements of the carpalia and tarsalia showed that the Natufian dogs were much smaller. What was the weight of the Natufian dogs? There is a high correlation (r2 =0·899) between P4 and body weight in recent Levantine wolves (Figure 20). Hence, if we impose the measurements of P4 of the Natufian dogs along this regression line, we may get a

90

E. Tchernov and F. F. Valla

close estimation of their weights. The range of weights estimated by this procedure for the Natufian dogs varies between 11 to 16·7 kg. (Figure 20). Figure 9 also shows that the two Natufian dogs from Hayonim Terrace, and the two dogs from Shukhba are significantly lighter in comparison with the recent wolves from Israel. The wolf is a species that follows Bergmann’s rule both in time and space (Davis, 1981; Dayan et al., 1990, 1992; Dayan, 1994) under colder and wetter climatic conditions (Tchernov, 1982; Horowitz, 1989; Baruch & Bottema, 1991). Therefore we expect that (Dayan, 1994) ‘‘. . . size reduction in the Natufian specimens is opposite to that expected both as a response to climatic conditions and a response to co-evolution with other competitors’’ (p. 639) (and see also Dayan et al., 1990, 1992). Therefore the size reduction of the Natufian dogs would have been much more drastic if we could have compared them with Natufian wild wolves (from which we do not have any record yet), being heavier than the recent population. Naturally wild wolves (Tchernov, 1984) are not expected to be found around Natufian villages. There is high correlation between the femur and body size as found in recent wolves. Therefore we estimated that the size decrease of the femur was proportional to body weight in the Natufian dogs, the sizes of which were found to be closely similar to the small wolf from the Sinai desert which weighs 12·3 kg. According with the size of the femur we may suggest that the weight of the Natufian dogs was indeed around 12 kg. The Saluki dog, however, shows, in relation to its body size (z 12 kg) extremely long feet. In this case there is obviously a disproportional increase in the size of the femur, as well as all other foot elements, apparently due to a secondary artificial selection for cursoriality.

Conclusions The Emergence of dogs as a result of unconscious selection The earliest record for genuine sedentism has been described from the Natufian period (Bar-Yosef, 1983; Braidwood, 1975; Perrot, 1961; Valla, 1987). The Natufian sites (especially the earliest ones) have been found in a relatively limited area in the southern Levant. When a new habitat was created by the Natufians within and around the primaeval villages (Tchernov, 1984, 1991, 1993), a confrontation between humans and other species came into an abrupt reality. From this moment we face severe competition among the species with similar or (partly) overlapping niches. Sedentary people and those highly hierarchical competitors which are basically omnivorous and catholic which found themselves co-occurring broadly in and around the anthropogenic sites, the result of which, during a relatively short period on an evolutionary

scale, was the development of indirect commensal relationships. The unique habitats that were created around permanent settlement sites encouraged certain animals to invade and rapidly colonize the newly opened anthropogenic niches. Not only did the unique habitats which were created around long lasting sites encouraged certain animals to invade and rapidly colonize this special habitat, but most of the colonizers became facultative or obligatory commensals, with extensive morphological and behavioural changes, and in a few cases, they underwent full speciation (Auffray et al., 1988, 1990; Tchernov, 1984, 1991, 1993). Wolves were part of this phenomenon (see more details in Davis, 1981; Davis & Valla, 1978; Dayan 1989; Tchernov & Kolska-Horwitz, 1991). At present there are only a few more cases of Natufian burials associated with dogs. All other dogs found in a Natufian context show a substantial diminution of their body size. Cases of differential size reduction are common (Hole et al., 1969; Tchernov & Horwitz, 1991). As for cattle, convincing figures for this phenomenon were demonstrated by Grigson (1989). For some Swiss and Danish prehistoric sites a ‘‘classic’’ demonstration of significant size differences between Neolithic pigs and cattle and their wild ancestors was published by Higham (1968). Davis (1981) illuminated a few clear-cut cases of size diminution during the end of the Pleistocene in the southern Levant, mainly in Capra and Sus, and compared them with recent populations. For the same region CluttonBrock (1979) has shown clear size differences between wild goats and pigs and their domesticated forms through the sequence of Jericho. That the phenomenon of size diminution in early domesticates of Capra, Ovis, and Bos was independently developed in Mehrgahr, Balucistan, was argued by Meadow (1984). Indeed, the course of size diminution in many regions reinforces our idea of the universality of body size diminution under anthropogenic domain (Tchernov & KolskaHorwitz, 1991). Several explanations have been offered to account for body size diminution with the advent of domestication. Some researchers have emphasized those features associated with methodological (conscious or direct) selection by humans for smaller animals (Darwin, 1875), being easier to control and more docile (Zeuner, 1963; Boessneck & Driesch, 1978; Meadow, 1984), to optimize grazing better (Jarman & Wilkinson, 1972) and to mature earlier. It also has been proposed that diminution resulted from selection for increased litter size rather than on body size or quality of the stock (Boessneck & Driesch, 1978; Rindos, 1984). In contrast to pressures resulting from direct human actions, unconscious or indirect selection pressures have been proposed as being responsible for smaller body size. Olsen (1985) stated that ‘‘large canids have mutually compatible social organization that eventually led to taming and, ultimately, to domestication’’ (p. xi). Yet it seems that there is no

Dogs from the Southern Levant 91

place to speak in terms of intentional domestication of wild wolves. It often has been claimed that many desired traits were intentionally (consciously or artificially) selected by humans during the process of domestication (Price, 1984; Rindos, 1984). Yet many of the traits, such as limiting agility, decreasing aggressiveness, shifting to an omnivorous diet and into more euryoecious habitats, more precociality, earlier maturity, size diminution (Price, 1984), may not be ascribed to artificial selection at all. Early domestication of animals may actually represent an intraspecific displacement along the K-/r-selection continuum, a shift that could have been affected by many other traits (Tchernov & Kolska-Horwitz, 1991). Although on the macro scale both wild and domestic terrestrial mammals are over all K-selectionists (Pianka, 1970, 1972; Southwood, 1981), the advent of domestication is characterized by an intraspecific shift to a more r-selected strategy along the general scale, although domestic ungulates are not r-selected strategists in the sense defined by Pianka (1970, 1972). Given a change in external conditions, a shift in strategy (from less r- to more r-selected, or from more K- to less K-selected) would have developed independent of human intervention. An excellent example is the variation in frequency of twinning in wild sheep. Geist (1971) has reported that twinning is very rare in North American bighorn and Dall sheep. In contrast, Valdez (1976) has reported exceptionally high frequencies of twinning in Asiatic Urial sheep. An additional example is the intraspecific variation in growth rates, timing, and duration of births in bighorn sheep living in low- as opposed to high-altitude environments (Risenhoover & Bailey, 1988). Under the low intraspecific competitor/predator environmental conditions offered by domestication, favourable morphobehavioural characters, such as r-selection strategies, were selected for out of the total range of variation exhibited by the species. Indeed, r-selection is considered an advantageous strategy for organisms colonizing new habitats (Safriel & Ritte, 1983). Natufian dogs are not different from other commensal animals (Auffray et al., 1988, 1990; Tchernov, 1984) which were attracted to the special and isolated habitats that were created within and around the sedentary human habitations, where they were at least quasi-isolated from wild wolves, and underwent rapid morphogenetic changes. Dogs (Davis & Valla, 1978), as well as other commensal animals (Mus musculus, Rattus rattus, Passer domesticus) (Tchernov, 1984, 1991), came into existence as new species (Auffray et al., 1988, 1990) during the Natufian period, and within the anthropogenic milieu. Hence we argue that any animal that is either deliberately or naturally forced or attracted to such an ecological regime will sooner or later shift to r-selection strategy, changing some behavioural and morphological traits accordingly.

The observed change in body size under domestication reflects a shift along the continuum from selection for individual viability toward local selection for higher reproductive rate, as a response to changed environmental conditions under the newly created habitats. It is proposed here that some, if not many, of the domestic traits that started to show up at the beginning of the ‘‘domestication’’ period could be by-products of the special environmental pressure induced on the animal populations around and within human habitations, and not the results of intentional selection imposed by people on their captive populations. That morphological changes during the process of early domestication (including dogs) are unlike to be direct product of intentional human selection has already been shown by Heiser (1988), Rindos (1984), Tchernov & Kolska-Horwitz (1991) and Zohary (1984), and specifically for dogs by Morey (1992). The theoretical importance of these changes is two-fold. First, their consistent occurrence indicates that general evolutionary process affected all early domestic canid populations similarly despite variability in cultural circumstances. Second, other domestic animals often exhibit similar changes (Clutton-Brock, 1981; Epstein, 1971; Zeuner, 1963), suggesting that this evolutionary process is of widespread importance under conditions of domestication (Wayne, 1986c). These two considerations suggest that emphasis on human selection as the primary agent responsible for changes in a domestic animal (Fox, 1978; Davis & Valla, 1978; CluttonBrock, 1984) may be misleading, at least for its earliest phases. As argued by Morey (1992: p. 82) ‘‘there is no warrant to assume that adopted pups were subjects to conscious, long-term domestication efforts’’ (see also Rindos, 1984; Tchernov & Kolska-Horwitz, 1991), ‘‘nor is it reasonable to suppose that different human populations all guided domestic canid evolution along the consistent path it initially took.’’ Evolutionary forces independent of human control do not necessarily cease simply because a domestic relationship is underway. Wayne (1986a) has pointed out that the cranial variables of modern breeds of dogs show negative allometry in relation to skull length. Therefore wild wolves show more developed post canine teeth in comparison with dogs, and explains it as due to intentional selection. We explained the changes of early domesticates, including dogs, as the result of ‘‘natural’’ changes under the special anthropogenic conditions around the early villages, when ‘‘artificial selection’’ (Wayne 1986a: p. 247) was not yet operated by Natufian people (Tchernov & Kolska-Horwitz, 1991; Morey, 1992). Wayne (1986a,b,c) deals with conscious domesticated breeds of dogs, mostly intentionally selected from special traits. When we deal with Natufian dogs, we believe that all the observed morphological differences between wild wolves and those early dogs are the outcome of unconscious selection, or rather the

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consequence of drastic change in the commensal populations under the new anthropogenic environment which were created at the beginning of sedentism in early Natufian period. Hence there is no way to speak about, at this early stage of ‘‘domestication’’ (or rather commensalization), of morphological changes under artificial selection. Morphological characteristics of the Natufian dogs During the Natufian period we witness an overall increase in body size of many species of mammals and birds (Davis, 1977, 1981; Kurtén, 1965; Tchernov, 1979), a phenomenon that has been described for many other places, and explained as a Bergmannian ecophysiological effect (Dayan, 1994; Dayan et al., 1991). The wolf is not exceptional (Davis 1981; Dayan 1994). It was hence expected that we would have found large-sized wolves in the Natufian deposits, but instead what we have are dog-like and dog-size animals which are even smaller than the arid form of Canis lupus pallipes from the Arabian desert (Dayan, 1994; Mendelssohn, 1982). Wayne (1986a,c) found that P3 and P4 lengths among modern dog breeds are characterized by negative allometry in relation to total skull length, while wild canids exhibit positive allometry. Indeed wolves display proportionally longer premolars than dogs of comparable size. Hence he deduced that tooth dwarfism at large body size reflects ‘‘artificial’’ versus ‘‘natural’’ selection (Wayne, 1986a: p. 247) in dogs versus wild canids. In general tooth size reduction was often held as an eventual correlate of canid domestication (Bökönyi, 1973, 1974, 1983; Clutton-Brock, 1970, 1984; Clutton-Brock et al., 1976; Scott, 1968). However, care must be exercised when describing tooth size changes. An alternative perspective is that body size has changed faster than tooth size in the evolving dogs. Morey (1992; p. 198) has pointed out that frequently ‘‘occurrence of conspicuously small teeth among large modern breeds of dogs distracts attention from the fact that smaller dogs often have proportionally longer teeth’’. He also showed that the pattern of ‘‘strong negative allometry for P3 and P4 lengths among modern dogs is anomalous in relation to allometric patterns among wild Canidae as a whole’’ (p. 198), but added that ‘‘a possible exception is the carnassial’’, which ‘‘yielded a relatively tight pattern of interspecific negative allometry in the genus Canis (including dogs)’’. Gingerich & Winkler (1979) and Pengilly (1984) argued that reduced variability is not surprising given the morphological and occlusal complexity of carnassial teeth in canids. There is a general consensus that relatively large and therefore crowded teeth are characteristic of the earliest domestic dogs (Bökönyi 1973, 1974; 1983; Clutton-Brock 1970, 1984). Gould (1975) pointed out that rapidly dwarfed lineages often exhibit relatively enlarged teeth, a pattern probably relating to lack of

tight developmental integration between dental growth and overall growth (Shea & Gomez, 1988). Among canids, lack of tight integration is indirectly suggested by consistently weak intraspecific correlations between tooth lengths and skull lengths (Wayne, 1986a). Do the Natufian dogs follow this scenario? The main region of the snout which underwent significant shortening was mainly restricted to the anterior portion of the mandible and the maxillary. But recent wolves and Natufian dogs show no difference in their width of the snout. This disproportional reduction of the snout included also a disproportional reduction in the size of the third premolars, but to a much less significant degree in the first and the second premolars. The canines are very conservative in their shape and size and indeed were only slightly changed in relation to many other variables. In relation to the length of the anterior part of the shortened snout it even became somewhat larger. This explains why the depth of the ramus in the canine region also remained relatively unchanged. M1 underwent a severe decrease in its size in spite of the fact that this region along the ramus became only slightly shorter. P2 and M1 also underwent a significant reduction in size, especially in comparison with the rear part of the maxilla. Consequently, as the reduction in the length of the muzzle included also some of the teeth, no crowding of the molars could have been found in the early Natufian dogs. Teeth crowding happened most probably in later periods when intentional, or directional selection was regularly practiced by people. M2 show minimum changes in size, but as the rear part of the mandible also did not change a lot, it remained proportionally the same, and no crowding could have been created. In many mandibular, maxillary and dental landmarks, if measured against a centroid size (like body size or weight) (Bookstein, 1991) shows a regression line typical to a relative growth rate. It is only the anterior region of the snout which shows a marked disproportional shortening when compared to wild wolves. Actually it can be regarded as a developmental retardation of the exterior part of the muzzle, a paedomorphosis, typical to other mammalian groups, like some primates. For the sake of occlusion it is expected that the maxillary and the mandibular teeth will have to change harmonically. Yet actually the teeth have changed their sizes independently and still kept the occlusion intact. Hence within a certain limit the lower and upper teeth may change their sizes disproportionally, and may explain the differential change in the lower and upper teeth. The dogs from Shukhba show the most extreme changes in comparison with wolves, and hence represent the most ‘‘doggish’’ form of all the known Natufian dogs in the southern Levant. The extreme variability in the size of the molars, especially in the fourth premolars and the first molars in the Natufian dogs indicate a progressive relaxation in natural

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selection pressure of these quasi-isolated populations of dogs. They were exposed to completely different kind of selection factors than wild wolves. The amount of similarity with wild wolves among these dogs possibly reflects the amount of gene flow between commensal and wild flocks. As for the post-cranial skeleton it seems that there is a proportional decrease in the length of the limb bones, while some of the epiphyseal regions, or the articular facets in some of the limb bones were disproportional changed, usually towards a lesser functional level, showing again a relaxation in the selection pressure of these populations. The lengths of the limb bones were not yet affected by conscious selection, and only show that these dogs did not need to run over large distances. The disproportional lengthening of the limb bones found in the Saluki dog is a typical example of intentional selection for cursoriality. In summary, although the Natufian dogs show a ‘‘classical characteristics of early domestication’’ (Dayan 1994: p. 637), ‘‘slight diminution of the carnassials alongside a marked reduction of mandibular and maxillary length’’, the shortening of the snout actually included mainly its anterior portion, while the posterior region remained only slightly affected, a common phenomenon in many long-snouted vertebrates (Tchernov, 1986). Moreover, the diminution of the carnassials were marked enough, at this earlier phase of their ‘‘domestication’’, that no crowding is displayed by any of the Natufian dogs. It is left to be shown that the reduction in the size of the snout of the Neolithic dogs from the southern Levant was marked enough to cause a crowding of their teeth. All this disproportional reduction of snout versus teeth, happened in later periods. As Dayan (1994) argued, the Neolithic dogs from Jericho show ‘‘still smaller carnassials in short mandible’’ (p. 637). We argue that this kind of morphology is typical to unconscious selection; no interference of people is displayed in the morphology of the Natufian dogs. This also explains the marked differences between the dogs from the different sites. Although the number of specimens of Natufian dogs is still small, it is already large enough to demonstrate the emergence of some patterns which indicate that all the recorded Natufian dogs are different even from the recent small-sized southern Levantine wolves and constitute a unique group, in spite of its wide morphological and size variability. Therefore Olsen’s (1985) chief criticism that southern Levantine early domesticated dogs are merely aberrant specimens is no longer justified.

Acknowledgments Our thanks are extended to ‘‘Sous-Direction des Sciences Sociale, Humaine et de l’Archéologie’’ of the ministry of foreign affairs (Paris) for financing the field work at Hayonim Terrace. We thank the ‘‘Centre de

Recherche Français de Jérusalem’’ for technical and administrative assistance, and the ‘‘Authority of Antiquity’’ for constant help and support. Special thanks go to O. Bar-Yosef for his help and advise; to B. Arensburg, L. Astruc, B. Boyd, R. Buxo, H. Colleuille, H. Khalaily, F. Le Mort and B. Poree for helping in the excavations. For assistance in the preparation of archeological and paleontological material we thank M. Chech. Special thanks are also extended to M. Barazany for her assistance in photography, to D. Ladiray for the drawings, and for polishing the English to B. Boyd.

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