osteology of thekurimaori: the prehistoric dog of new zealand

Journal of Archaeological Science (1997) 24, 113–126

Osteology of the Kuri Maori: the Prehistoric Dog ofNew Zealand

Geoffrey Clark

Division of Archaeology and Natural History, Australian National University, Canberra, ACT 0200, Australia

(Received 4 July 1995, revised manuscript accepted 5 November 1995)

This paper describes osteometric dimensions and anatomical features of the Polynesian dog of New Zealand (kuri) andphenotype reconstruction is made using regression formulae. Compared to other prehistoric dog populations, kuri werefound to exhibit low to moderate levels of morphological variation. Studies of ancient dog remains have tended to focuson the metric variation found in cranial remains, and post-cranial elements are not reported. It is suggested that aholistic approach, incorporating anatomical and osteometric data from cranial and appendicular remains, will providebetter archaeological data on which to base population comparisons. ? 1997 Academic Press Limited

Keywords: KURI, DOG, PREHISTORIC, OSTEOLOGY, NEW ZEALAND, POLYNESIA.

Introduction

T he kuri was the single domesticated animal keptby Maori and the only terrestrial mammaliancarnivore introduced during New Zealand’s

prehistory. Thus, the kuri had a unique relationshipwith Maori and place in New Zealand’s pre-Europeanecology. Following European contact a number ofdescriptions of this animal were recorded, the mostdetailed being that of Crozet (1891: 76), who observedkuri in 1772:

‘‘The dogs are a sort of domesticated fox, quite black orwhite, very low on the legs, straight ears, thick tail, longbody, full jaws but more pointed than that of the fox, anduttering the same cry; they do not bark like our dogs.These animals are only fed on fish, and it appears that thesavages only raise them for food.’’

However, no dogs were collected during the earlycontact period, although a few mounted specimens,dating to the late 19th century, are now regarded astype specimens of the breed (Figure 1). As the kuri isextinct, it will be primarily through the analysis of itsskeletal remains that the kuri, its affinities to otherPacific dog populations and its relationship with Maoriwill be understood.This paper is based on the analysis of 1369 cranial

and post-cranial remains. Assemblages of kuri bonesfrom 38 New Zealand archaeological sites, rangingfrom Mason Bay in the extreme south to Tom BowlingBay in the far north, were examined (Appendix 1).The antiquity of the remains spans the whole ofNew Zealand’s prehistory (Anderson, 1991). Over 9500measurements were taken, along with a number ofdental and non-metric observations. The study was

110305–4403/97/020113+14 $25.00/0/as950098

undertaken with two major aims in view: firstly, toestablish a comprehensive osteometric and osteologicaldescription of the kuri; secondly, to establish the degreeof population variability which existed during theprehistoric period.The results provide the metric population par-

ameters of the Polynesian dog in New Zealand, againstwhich tropical Polynesian assemblages can be com-pared. It is worth noting that despite the relativeubiquity of commensal remains in Polynesia (Kirch,1984: 88), little osteological analysis has yet taken place(Kirch & Weisler, 1994: 295). The benefits of suchstudy in elucidating prehistoric migration patternshave, however, long been recognized (Wood-Jones,1929; Luomala, 1960: 222; Groves, 1983: 119).

Previous StudiesOf the commensals—the pig (Sus scrofa), the dog(Canis familiaris) and the fowl (Gallus gallus)—introduced into Oceania by prehistoric Polynesians,only the dog was established successfully in NewZealand. The absence of the pig and fowl is intriguingconsidering the large size of New Zealand in relationto most Polynesian archipelagoes and the abundantresources which existed during colonization. A failureof these animals to survive the long voyage fromcentral East Polynesia (Bay-Petersen, 1984), a deliber-ate decision not to import these animals, based onreports brought back by the first discoverers of a largeand plentiful fauna (McGlone et al., 1994), or a limitedintroduction, followed by extirpation once the extentof wild resources was recognized (Flannery, 1995)appear to be the most likely explanations.

3? 1997 Academic Press Limited

114 G. Clark

The importance to Maori of the single domesticate—kuri—is shown by its depiction in rock art (Anderson,1981: 17; Bain, 1985), the use of its skins for statusgarment manufacture (Black, 1922; Mead, 1969: 241)and the presence of its remains in numerous NewZealand prehistoric sites (Allo, 1971: 30–31).Archaeological studies of kuri remains have estab-

lished its nutritive importance to the Maori economy(Allo, 1972; Shawcross, 1972; Smith, 1981a), theindustrial uses of its skeletal remains (Fisher, 1934;Duff, 1956: 95; Coutts & Jurisich, 1975), and, throughcoprolite and midden studies, its probable diet(Scarlett, 1972: 23; Byrne, 1973; Williams, 1980;Taylor, 1984: 221; Nichol, 1988: 330). A number ofresearchers have analysed kuri material for congenitaldental abnormalities (Teal, 1975; Smith, 1981b), fol-lowing the demonstration that dental variation mighthave a genetic basis (Allo, 1971). In some cases this hasenabled statements to be made about contact betweenprehistoric groups (Leach, 1979: 92–93; Davies, 1980:81–91).Initial osteometric and anatomical details of the kuri

skeleton were furnished by Haast (1872: 98), Hector(1877) and Hutton (1898). A major study of the kuri byAllo (1970) included limited metric information fromthe cranium and mandible, while the major limb boneswere only described using one measurement—that ofgreatest length (ibid: 66). A number of cranial indicesand anatomical observations of the skull and mandiblewere thought to be useful in identifying whether re-mains belonged to a kuri or were from a European

breed (Allo, 1970: 46–64, 1971: 31–35, Allo Bay-Petersen, 1979: 177–178). The list of features was aug-mented by Watt (1975: 141), who recorded that the kurimandible was characterized by a forward extension ofthe antero-lateral border of the ascending ramus.

Materials and Methods

Skeletal elements examined in this study were thecranium, mandible, scapula, humerus, radius, ulna,pelvis and tibia. Classification of the remains to one oftwo osteological categories (mature or immature) wasmade, using criteria developed by the author for thecranial remains (Clark, 1995) and by Sumner-Smith(1966) for the appendicular skeleton. Discussion here islimited to the data from osteologically mature remains.As most skeletal growth is achieved during early devel-opment (Hildebrand, 1974: 179), it is necessary toidentify skeletally mature from immature remains.Without this fundamental assessment it is impossibleto distinguish the component of variation caused bygrowth from variation due to genetic or environmentalfactors. Once adult stature is reached the degree ofbone morphology alteration is reduced and dimensionsare unlikely to change considerably (Wayne, 1986: 387;Reitz & Ruff, 1994: 699).Determination of sex from canine remains which are

fragmentary and disarticulated, as is the case for thekuri, is only possible using the cranium (The & Trouth,1976; Trouth et al., 1977). There is no established

Figure 1. Mounted example of a dog often regarded as the type specimen of the kuri breed.

Osteology of the Kuri Maori 115

method for assigning isolated dog remains to a particu-lar individual of a particular sex (Brothwell et al., 1979:145) and no direct attempt has been made to controlfor this component of variation.A total of 81 metric variables—32 cranial, 14 man-

dibular and 35 post-cranial—were recorded. Twostandard cranial indices—the cranic (cephalic) indexand the palatal index—were calculated (Shigeharaet al., 1987: 22). Description of the chord variables andindices can be found in Appendix 2, while the inter-ested reader is referred to von den Driesch (1976) andClark (1995: 280–297) for detailed definitions. Metricdata was recorded with digital callipers (Mitutoyo),accurate to 0·01 mm. Measurement error was assessedusing a procedure similar to that used in humanosteology (White & Folkens, 1991: 292). Percentageerror results were generally between 1% and 3% andwill not be discussed further. Descriptive statistics(sample number, mean, standard deviation, minimumand maximum values and the coefficient of variation,Cv) were calculated for each variable using standardformulae. All calculations were made using SPSSversion 4·0 for the Macintosh.Dental and non-metric cranial traits identified by

Allo as distinctive to the kuri (1970: 46–64, 1971: 35)were recorded. The features are: skull of long andnarrow proportions (dolichocranic); presence of aprominent sagittal crest; posterior end of the nasals(nasion) ending level with the fronto-maxillary suture(nasion is incorrectly described by Allo (1971: 32, 1979:177) as ending level with the premaxilla); presence ofsupernumerary alveoli (crania and mandibles) andwell-spaced dentition. The distribution of these traits inkuri cranial remains was compared to their presencein a modern dog sample of 52 crania and 90 demi-mandibles (Biochemistry Department, University ofOtago).Information for skeletal elements is grouped into

four categories: cranium, mandible, forelimb (scapula,

humerus, radius and ulna) and hind limb (pelvis, femurand tibia). As the dog is the most morphologicallyvaried animal on earth (Wayne, 1986: 382), it is likelythat the kuri will overlap in many of its anatomicalfeatures with modern breeds and cross-breeds (Allo,1971: 30–31). Therefore, looking for distinctive fea-tures that will separate kuri remains from those ofmodern dogs is unlikely to provide information ofarchaeological utility. The approach used here to dis-cuss kuri osteology is to define the metrical parametersof the skeleton and combine this with observationsdetailing the common anatomical features of the popu-lation, notwithstanding the occasional use of skeletalcomparisons that have been made on modern dogremains by various authorities.

The CraniumLateral view (Figure 2)—The mean length of 20 osteo-logically mature kuri crania is 170·6 mm (Table 1),similar to that recorded (167 mm) from a sample of 14

Figure 2. Lateral view of the kuri cranium (all scale bars in cm).

Table 1. Kuri cranis descriptive statistics (measurements in mm)

N Mean S.D. Min. Max. Cv

Length variablesSkull L. 20 170·6 10·4 154·7 191·4 6·1Condylobasal L. 20 156·4 10·2 138·9 178·2 6·5Facial L. 46 95·3 5·7 84·8 106·7 6·0Palate L. 44 85·1 4·4 76·0 94·8 5·2Neurocranium L. 18 81·4 5·7 74·1 92·1 7·0Snout L. 46 78·5 5·1 70·1 89·3 6·5Nasal L. 32 56·2 4·4 49·0 66·3 7·9

Breadth variablesBizygomatic B. 14 95·7 6·8 77·7 106·2 7·1Mastoid B. 20 62·2 3·5 56·2 70·9 5·6Palate B. 42 58·1 3·9 51·9 65·6 6·7Braincase B. 18 51·3 2·1 48·3 56·2 4·1Frontal B. 37 50·5 4·2 43·8 60·5 8·3Paraoccipital B. 21 45·1 2·4 40·9 49·6 5·3Postorbital B. 44 32·3 1·7 27·9 37·1 5·3Condyle B. 22 32·4 2·2 28·7 37·5 6·8Interorbital B. 43 31·9 2·3 28·3 37·3 7·2Min. Palate B. 46 31·7 2·2 28·1 35·8 6·9Bulla B. 22 18·6 1·6 15·6 21·2 8·6

Height variablesSkull H. 22 56·1 4·7 49·7 65·8 8·4Parietal H. 22 50·3 2·6 45·2 55·2 5·2Akrokranion H. 21 45·8 3·9 40·1 53·9 8·5Orbital H. 31 28·8 2·5 24·0 34·7 8·7Foramen H. 22 13·3 1·3 11·9 16·4 9·8

Dental variablesPremolar L. 49 39·3 2·3 35·0 45·4 5·9Molar L. 49 16·5 1·5 12·3 20·0 9·1P4 B. 46 8·3 0·5 7·4 9·6 6·0P4 L. 47 14·3 1·0 12·2 16·6 7·0M1 B. 46 11·7 1·0 7·4 13·4 8·5M1 L. 46 9·0 0·7 7·5 11·2 7·8M2 B. 31 8·2 0·6 7·2 9·6 7·3M2 L. 30 5·5 0·5 4·7 7·1 9·1

Cranial indicesCranic (cephalic) 12 56·8 1·6 54·0 59·4 2·8Palatal 35 68·5 2·4 64·2 74·1 3·5

116 G. Clark

crania by Allo (1970: 66). In lateral view the prominentdevelopment of the interparietal and nuchal crest canbe seen. The mean height of the sagittal crest above thedorsal parietal surface is 5·8 mm (Skull H. mean minusParietal H. mean). The facial region is slightly tomoderately dished and the low coefficient of variationfor this measurement (Facial L., Cv=6·0%) indicates asmall amount of variability in facial length. Miller et al.(1968: 8) note that in modern dog breeds the greatestvariation in skull shape occurs in the facial region. Thezygomatic arch and mandibular fossa, which receivesthe condyle of the mandible to form the temporo-mandibular joint, is well developed in mature dogs,indicating that kuri had a powerful jaw. The canine issmall and slender and the carnassial and molar teethare also small in comparison to the teeth of modernmedium-sized cross-breeds.

Dorsal view (Figure 3)

The mean bizygomatic breadth is 95·7 mm and has aCv of 7·1%. The external sagittal crest often reachesfrom the external occipital protuberance to beyond thefronto-parietal suture. The surface of the parietals isroughened in some specimens from attachment of thetemporalis muscle. The zygomatic process of the fron-tal bone is pointed and the shape is age-related—inolder animals the area becomes sharply pointed andpitted due to the action of the orbital ligament. Asnoted previously by Allo (Allo Bay-Petersen, 1979:178), the muzzle narrows at the third premolar to forma relatively short and blunt snout.

Basal view (Figure 4)

The posterior border of the palate is behind M2 for allof the kurimaxilla examined in this study. The positionand size of the palatine fissure, the anterior palatine

foramen and the palatine groove are highly variable.The basilar part of the occipital bone bears strongmarkings from insertion of the major and minor bilat-eral rectus capitas ventralis muscles. The tympanicbulla is compressed, with occasional ribbing aroundthe border of the auditory tube. Foramen (posterioralar, oval and hypoglossal) are frequently asymmetric.The intercondyloid incisure varies from a triangular toa half-rounded form.

Occipital view (Figure 5)

The major features of the posterior region of the kuricrania are the sagittal crest, dorsal nuchal line andforamen magnum. The dorsal nuchal crest rises fromthe mastoid process to form the external occipitalprotuberance. The morphology of the occipital protu-berance varies from a sharply pointed triangular formto a rectangular, flattened shape. The difference is

Figure 3. Dorsal view of the kuri cranium.
Figure 4. Basal view of the kuri cranium.
Figure 5. Occipital view of kuri crania.


related to the development of the interparietal process.As the interparietal bone develops from between theexternal sagittal crest, the dorsal nuchal region appearsto be wide and flat. Through time or because of a highdegree of muscle activity it fuses with the parietals andbecomes steeper and thinner due to the action of thetemporalis muscle, eventually forming a pointed, trian-gular shape in nuchal view. Little variation in the shapeof the foramen magnum was observed (Figure 5),despite the presence of a number of forms found inmodern dogs (Miller et al., 1968: 13; Shigehara et al.,1987: 21).A tighter range for the cranic index was found in this

study (54–59·4) compared to that found previously(51–63) and this might indicate that some of the re-mains examined by Allo were not kuri. Certainly, of the11 crania used by Allo to establish this index, bothunlocalized surface finds and juvenile crania were in-cluded (Allo, 1971: 31–32). The cranial index systemdevised by Miller et al. (1968: 8, figure 1–37), where thecranial index for brachycranic dogs=81, for mesati-cranic dogs=52 and for dolichocranic dogs=39, firmlyplaces the kuri crania as a mesaticranic form (head withmedium proportions), with a slight tendency towardsthe brachycranic skull shape (short and wide-headed).The number and type of dental abnormalities will be

treated in a separate paper. It is enough to note herethat 21% of the 120 maxilla examined with intact toothrows have some sort of dental abnormality. Of these,14% display supernumerary dentition, with an extra P1

occurring in 10% of all maxilla. In comparison, 16% ofthe modern dog crania have supernumerary dentalabnormalities, of which 8% involve P1.Almost 80% of all kuri crania examined (35 out of

44) have nasals ending level with the fronto-maxillarysuture and 20% have nasion ending posterior to thisposition. Of the 52 mesaticranic modern skulls, 67%have a nasion point level with the fronto-maxillarysuture.

The MandibleThe greatest mean length of the osteologically maturekuri mandible is 129·2 mm (Table 2). Coefficients ofvariation are between 4·9 and 5·2% for the threevariables describing mandible length (Mandible L.1–3). The ramus or vertical part of the mandible has alarge masseteric fossa for insertion of the massetermuscle. The coronoid process overhangs the condyloidprocess, as is the case for most domestic dogs. Situatedbetween the middle and posterior mental foramenthere is often one or more extra foramen. The shape ofthe angular process is age-related. In younger dogs it issmooth and rounded, while in older dogs the process iscontoured from muscle insertion (Figure 6).The greatest variation in the mandible measurements

is found in the height of the mandible body at M1(Corpus M1 H., Cv=8·3%) and the length of thepremolar row (Premolar L., Cv=8·3%).

The spacing between teeth in the premolar row ishighly variable. In some cases teeth are packedclosely together and in others they are well-spaced. Itis often difficult with fragmentary remains to distin-guish between developmental tooth-crowding, causedby the incomplete growth of an animal’s dentition,and congenital tooth-crowding. Overall, though, thereare few cases of restricted premolar tooth rows.Of 181 mandibular tooth rows, 39% have a patho-

logical or congenital abnormality. Supernumerary al-veoli contribute 22% of this total, with an extra M3(9%) and M3 double rooted or extra (7%) occurring themost frequently (when M3 is absent it is difficult to tellif an extra tooth or extra tooth root is present). Incomparison to the crania, where the frequency of extraalveoli is similar in kuri and modern dogs, the mandi-ble shows striking differences between groups. Themodern sample of 90 mandibles has a high incidenceof abnormalities (41%). However, only 6% involved

Figure 6. Lateral view of kuri (left) mandibles.

Table 2. Kuri mandible descriptive statistics (measurements in mm)

Mandiblevariables N Mean S.D. Min. Max. Cv

Mandible L.1 106 129·2 6·6 117·5 145·1 5·1Mandible L.2 115 128·4 6·7 118·2 145·1 5·2Mandible L.3 119 123·2 6·1 112·8 138·5 4·9M3-P1 L. 118 64·2 3·2 56·3 72·5 5·0M3-P2 L. 118 58·1 2·8 49·7 64·5 4·8Ramus H. 86 54·8 4·1 45·9 64·3 7·5Corpus M1 H. 118 22·8 1·9 17·4 28·2 8·3Corpus P2 H. 117 18·5 1·4 14·6 27·7 7·6Premolar L. 117 35·1 2·9 24·8 53·7 8·3Molar L. 117 29·7 1·9 23·0 35·4 6·4M1 B. 96 7·3 0·4 6·2 8·4 5·5M1 L. 96 17·1 0·9 15·0 19·3 5·3M2 B. 65 5·7 0·4 4·2 6·8 7·0M2 L. 65 7·4 0·6 6·2 8·9 8·1

118 G. Clark

supernumerary alveoli and 2% of these occurred onthe M3.

Forelimb Elements

Scapula

The mean height along the spine of the kuri scapula is99·3 mm (Table 3). This variable has a high Cv value of13·3% and a frequency graph shows that the data isnegatively skewed. This distribution is almost certainlythe result of the earlier development of the scapula(Sumner-Smith, 1966: 30), which fuses in the period ofrapid growth (in modern dogs between 4–5 months),rather than in a period of slowed growth (>6–7months).The caudal border of the scapula is straight and

thickened. The spine is prominent and in some speci-mens rises to around 2 cm in height. The acromion iswell developed and is rolled posteriorly. The positionof the two nutrient foramen, located on the lateralsurface at the base of the spine and on the medialsurface of the caudal border, appears to be stable. Inrelation to modern medium-sized breeds, the cranialborder of the kuri scapula is rounded and the dorsalfourth of the caudal border, which serves for muscleattachment, is well developed (Figure 7). The medialsurface is characterized by muscular lines caused byinsertion of the m. subscapularis. The shape of the kuriglenoid fossa is not especially different from that ofmodern dogs, despite Allo’s assertion that it was longerand narrower (1971: 35).

HumerusThe greatest mean length of the kuri humerus is122·5 mm (Table 3). The coefficient of variation for thelength variable is 5·6% which is the smallest Cv for thiselement. All variables recording aspects of humerusbreadth and depth have Cvs approaching, and in onecase (Midshaft (a–p)) exceeding, 10%.The proximal portion of the body and neck of the

kuri humerus has a pronounced twist. This can be seenin Figure 8, where three kuri humeri are compared tothose from three modern cross-breeds. When placedwith their caudal surface resting on a flat surface themodern dog humeri are stable, resting on the caudalsurface of the humeral condyle distally and on thehumeral head proximally. The twist in the proximalportion of the kuri humerus causes it to roll so that it

Figure 7. Lateral view of kuri (left & right) scapulae.

Figure 8. Kuri and modern dog (right) humeri showing the twist inthe kuri corpus.

Table 3. Kuri forelimb descriptive statistics (measurements in mm)


Scapula variablesScapula L. 36 99·3 13·2 62·4 116·2 13·3Glenoid L. 34 22·7 1·8 17·7 25·9 7·9Glenoid B. 32 13·8 1·8 10·7 22·0 13·0

Humerus variablesHumerus L. 32 122·5 6·9 108·7 137·0 5·6Proximal D. 42 30·9 3·0 25·9 37·7 9·7Midshaft (m–l) 32 11·1 1·0 8·9 13·1 9·0Midshaft (a–p) 32 16·4 1·8 12·1 20·4 11·0Distal B. 29 28·3 2·4 24·7 32·6 8·5

Radius variablesRadius L. 35 114·5 8·2 95·1 143·2 7·2Proximal B. 32 16·1 2·0 13·2 20·2 12·4Proximal D. 33 10·2 1·2 8·6 12·9 11·8Midshaft (m–l) 35 12·7 1·7 9·6 16·5 13·4Midshaft (a–p) 35 8·1 1·4 5·7 11·7 17·2Min. shaft (m–l) 45 11·5 1·6 8·7 15·6 13·9Distal B. 53 19·0 2·4 15·7 25·1 12·6Distal D. 51 11·2 1·4 8·9 15·3 12·5

Ulna variablesUlna L. 27 134·5 8·5 111·6 151·3 6·3Olecranon D. 27 17·6 1·6 14·7 21·5 9·1Anconaeus D. 27 21·9 2·3 17·2 26·7 10·5


lies with its lateral surface facing the ground surface.Although the causes of the twist in the kuri humeribody and neck are probably complex, as is the casewith the shape of the Polynesian human femur(Houghton, 1980: 35), it appears that the prominentdevelopment of the anconeal crest is partially respon-sible. The muscles attaching to the anconeal crest arethe m. brachialis and caput lateral of the m. triceps.The action of these muscles is in flexion of the elbowjoint. The medial and lateral epicondyles of the distalepiphysis are often pitted, with occasional nodularbone formations (arthropathy).The radial and olecranon fossa communicate with

each other by the supratrochlear foramen (STF). Thepresence of an open or closed STF has been recordedfor all distal humeri, as it had been suggested that theabsence of a STF might be genetically determined(Allo, 1970: 87). Of 32 mature humeri, 59% had anopen STF, while only 19% (N=16) of immature humerihad this feature. It is likely, then, that rather than agenetic cause, there is a correlation between the pres-ence of an open or closed STF and a dog’s skeletaldevelopment. This is also the case with the presenceof an open or closed STF in the human humerus(Wolpoff, 1980: 32).

Radius

The least amount of variation was found in the radiilength measurement (mean 114·5 mm, Cv=7·2%). Theproximal head (caput radii) and distal extremitymeasurements have Cvs between 12 and 13%, with thegreatest variation occurring in the shaft diameter vari-ables (Table 3). For example, midshaft diameter (a–p)has the highest Cv value, at 17·2%. This high Cv valueseems partly the result of the development of theinterosseous crest in some individuals. Muscles whichattach in the interosseous crest line are extensorsinvolved in the extension and rotation of the forelimb.A common feature of the kuri radius is the highly

developed eminence for the attachment of the cranialcrus of the lateral ligament of the elbow joint (Figure9). The nutrient foramen in most kuri radii is large andtends to be located in the distal section of the rough-ened area, caused by attachment of the interosseousmembrane, which, in life, unites the radius to the ulna.The medial articular circumference of the radius headis often stepped, with occasional arthropathic forma-tions (see below). A feature of the kuri radius is thedegree of curvature of the corpus compared to modernbreeds.

Ulna

The mean length of 27 complete kuri ulna is 134·5 mm(Table 3). This is slightly smaller than the result foundby Allo (1970: 71), who recorded a mean length of137 mm, using a sample of 18 ulna. The greatest

amount of variation was found in the depth of theprocess anconaeus (Anconaeus D., Cv=10·5%).The distal shaft of the kuri ulna is curved posteriorly

in comparison with some modern breeds. The medialand lateral borders of the trochlea notch are oftenpitted (Figure 10). In some specimens the pitting hasmodified much of the anconeal process. The radialnotch also has evidence of this condition. Bone pittinghas previously been identified as indicating osteoarthri-tis in kuri (Allo, 1970). It is perhaps more correctlytermed arthropathy to distinguish it from ‘‘arthritis’’which is a term usually reserved for degenerativediseases of infectious origin. The term arthropathyrefers to joint changes which are non-infectious. Indomestic animals arthropathic formations have beenlinked in archaeological sites to the presence of old oroverworked animals (Siegal, 1976: 362).

Hind Limb Elements

PelvisThe mean length of 29 kuri pelvi is 118·2 mm (Table 4).The coefficient of variation is 10·9% which mightindicate that the pelvis forms slightly earlier than thelong bones and therefore encompasses a greater degreeof developmental variation. Some support for this ideais given by the frequency distribution of this variablewhich shows that the data is negatively skewed. Thereis, however, little information on the time of completefusion of the main pelvic bones, in comparison to thatof the limb bones (Hare, 1960; Sumner-Smith, 1966;Adams, 1986).Average length of the acetabulum is 18·6 mm,

slightly under the 20 mm that Miller et al. (1968: 78)found for the average length of medium-sized dogs.The greatest amount of variation found in the pelvis

Figure 9. Caudal view of kuri (right) radii.

120 G. Clark

measurements is in the ilium body dimensions (IliumH. and Ilium B.), which have Cvs of 11·0% and 13·3%respectively. The kuri pelvis has a dorsal iliac spinewhich is well developed and in average specimens

approximately 1 cm across at its widest point. Milleret al. (ibid: 80) comment that the iliac spine in largeworking breeds reaches a width of nearly 1 cm. Theiliopectineal eminence is also well developed. Thearticular surface of the pelvis is rugged, while the iliumwing is short and broad. Variation in the shape of theilium wing was noted (Figure 11). Whether this mor-phology represents a sexual dimorphic feature is uncer-tain, although some aspects of the complete pelvis arethought to be of use in differentiating male from female(Sisson, 1930: 202; Shigehara et al., 1987: 21).

FemurThe mean length of the kuri femur is 137·2 mm—thegreatest mean length of any kuri limb bone (Table 4).In modern breeds and cross-breeds the ulna tends to bethe largest bone (Miller et al., 1968: 72). A similarmean length of 139·3 mm was found using a sample ofnine femora (Allo, 1970: 71). Coefficients of variationare under 10% for most variables. One midshaft diam-eter (m–l) and the depth of the caput femoris have Cvsof 11·1% and 9·9% respectively.The kuri femur has a greater trochanter which is

either level or slightly higher than the most proximalpoint of the femur head (Figure 12). Sisson (1930: 20)comments that the greater trochanter of modern dogsdoes not extend as high as the femur head, while Milleret al. (1968: 84) note that the greater trochanterextends almost to the frontal plane in modern dogbreeds.The greater, lesser and third trochanter are well

developed, as are the distal, medial and lateral bordersof the popliteal surface. The shaft is straighter than theshafts of the forelimb bones, which are noticeablycurved in the midshaft region. The fovea on the head issmall and often difficult to locate except as a slightly

Figure 10. Medial view of kuri (right) ulnae. Note arthropathicformations on the trochlea notch.

Figure 11. Lateral view of kuri (right) pelvi.

Table 4. Kuri hind limb descriptive statistics (measurements in mm)


PelvisPelvis L. 29 118·2 12·9 80·4 133·7 10·9Acetabulum B. 29 18·6 1·7 15·7 22·1 9·1Ilium H. 29 16·4 1·8 13·4 20·6 11·0Ilium B. 29 9·0 1·2 6·7 12·2 13·3

FemurFemur L. 23 137·2 10·2 115·9 160·1 7·4Proximal B. 43 32·2 2·1 27·9 37·6 6·5Caput D. 51 15·2 1·5 10·1 18·3 9·9Midshaft (m–l) 32 11·7 1·3 9·4 14·6 11·1Midshaft (a–p) 31 12·2 1·2 10·0 15·4 9·8Distal B. 23 27·3 2·3 22·9 31·8 8·4

TibiaTibia L. 30 128·3 9·2 107·4 146·9 7·2Proximal B. 31 28·9 2·4 25·6 33·6 8·5Midshaft (m–l) 30 11·3 1·3 8·6 13·9 11·5Midshaft (a–p) 30 12·5 1·4 9·4 14·8 11·2Distal B. 30 18·3 1·5 22·4 22·4 8·2Distal D. 30 14·0 1·3 17·0 17·0 9·3


roughened area. The position of the nutrient foramenon the caudal surface of the shaft is highly variable—itis usually found on the trochanteric surface but canoccur in the midshaft area.

Tibia

The mean length of the kuri tibia is 128·3 mm (Table4), slightly smaller than that recorded by Allo(131·3 mm) using a sample of 14 tibia. The tibial lengthCv is the smallest of any variable, at 7·2%. Coefficientsof variation are between 8 and 9% for proximal anddistal epiphysial measurements. The highest Cv valuesare found in the midshaft diameter variables (m–l,Cv=11·5%, a–p, Cv=11·2%). There appear to be rela-tively few distinguishing characteristics of the kuritibia. The shaft is straight and the position of thenutrient foramen appears to be relatively stable (Figure13). This foramen is found on the lateral edge of thecaudal surface, approximately half-way between themidpoint of the shaft and the proximal epiphysis.

Phenotype ReconstructionEarly European accounts of the kuri noted smalloverall size, with short legs in relation to a long body,variable coat colour, a broad head with a pointedsnout and the absence of a bark (Forster, 1778: 189;Crozet, 1891: 76). The breed’s dimensions were neverrecorded and since 1830 kuri genes have been absorbedinto the gene pool of European dogs (Colenso, 1878).In the absence of any first-hand quantification ofdimensions, regression formulae based on skeletal di-mensions has been used to estimate some phenotypeparameters. Shoulder height is calculated using the

equations developed by Harcourt (1974: 154), whilemethods for determining body weight from the corpusheight of the mandible and the minimum circumfer-ence of the femur follow those of Wing (1978),Hamblin (1984) and Anderson et al. (1985).The average shoulder height of the prehistoric kuri

is just under 40 cm (Table 5), indicating a small tomedium sized animal—slightly bigger than a cockerspaniel but smaller than most border collies. Weightestimates using three different formulae give similarresults, suggesting a mean weight of 13–15 kg. Incomparison to 15 modern breeds, kuri mean weightis greater than that of spaniels but less than that

Figure 12. Caudal view of kuri (right) femora.
Figure 13. Lateral view of kuri (right) tibiae.
Table 5. Kuri shoulder height (cm) and body weight (kg) estimates

N Mean Min. Max.

Height estimation (cm),after Harcourt (1974)Humerus (3·43#G.L.)"26·54/10 32 39 35 44Radius (3·18#G.L.)+19·51/10 35 38 32 47Ulna (2·78#G.L.)+6·21/10 27 38 34 43Femur (3·14#G.L.)"12·96/10 23 42 35 49Tibia (2·92#G.L.)"9·41/10 30 38 32 44Kuri mean shoulder height (cm) 39 34 46

Weight estimation (kg)Mandible height at M1 (mm) 118 22·8 17·4 28·21. Weight (kg) 118 13·8 7·8 21·62. Weight (kg) 118 15·2 8·3 24·6Femur circumference (mm) 9 42 35 493. Weight (kg) 9 14·2 8·1 21·1

1. Log y=2·1122 (log #)+1·2722, Log y=weight (g), Log#=mandible height at M1 (mm), after Hamblin (1984).2. Log y=2·2574 (log#)+1·1164, after Wing (1978).3. Weight (g)=0·35·femur circumference (mm)/1·50, after Andersonet al. (1985).

122 G. Clark

of bulldogs and border collies (Kirk, 1966). Skeletalobservations which provide further insight into thephenotype of this animal include the presence of asharply pointed snout and curved forelimbs witharthropathic formations occurring on the joints. Theyindicate why the kuri was noted as having a fox-likeface (Crozet, 1891: 76) and suggest that the animalhad heavy forequarters—a trait noted in rock artrepresentations of the kuri (Anderson, 1981).

DiscussionOver the past 120 years knowledge of the kuri has notappreciably increased, due largely to the reliance ofprehistorians on observations collected from the his-torical records (Colenso, 1878; White, 1894; Thomson,1922; Luomala, 1960; Titcomb, 1969; McCulloch,1986). Few of these have attempted to incorporatearchaeological or osteological data into their discus-sion (see Anderson (1990) for an exception).This paper has presented osteometric dimensions for

the major skeletal elements and described some of thecommon anatomical landmarks of the kuri skeleton.Anatomical features, particularly those involving thecranium, have in the past been used to identify whethera canid was kuri or of European origins. While some ofthese features are relatively ubiquitous, such as thepresence of a prominent sagittal crest and the positionof nasion in relation to the fronto-maxillary suture,they are also found in reasonable frequencies in theskulls of modern dogs. When canine remains are ofuncertain affinities, such as those from historic Maorisites, it is appropriate to use metric data as well asobservations of anatomical features to classify theremains.Reassessment of the kuri crania as mesaticranic

rather than dolichocranic means that supernumeraryalveoli can no longer be considered a reliable kuri/European dog separator. It is worth noting that extraalveoli involving the mandibular M3 do seem to occurwith greater frequency in the Maori dog and this traitmay have some diagnostic utility. Although it has beenassumed that such alveoli are inherited, Wolsan (1984)found that diet, trauma and infection can lead tosupernumerary dentition through splitting of the devel-oping tooth bud. More work needs to be undertaken toclarify the causes of the various abnormalities beforegenetic links between prehistoric Maori groups can beinferred on the basis of shared dental abnormalitiesfound in dogs from different sites.Limited phenotype reconstruction using regression

formulae allows the quantification of body parametersand comparison with descriptions given in the earlyhistorical records. There are many descriptions of dogswhich are claimed to be true kuri in the period 1830–1890 (Luomala, 1960: 220), but by this time Europeandogs had become established and it is difficult todetermine whether kuri, kuri-European crosses orEuropean dogs are being described.

Population variation

The degree of skeletal variation present during theprehistoric period was assessed using the coefficient ofvariation, a frequently used statistic for examiningpopulation variation. The Cv is a useful measure, asthere is a general tendency for morphological varianceto scale proportional to the square of the variablemean (Lande, 1977: 214). Although there is no Cvfigure that can be used to indicate whether prehistoricmodification of kuri took place, studies of other speciessuggest that values over 10% should be carefullyscrutinised (Groves, 1991: 210).Studies of prehistoric dog remains from other parts

of the world display a range of Cv figures (Cvsrecalculated from original data). Low values found inJomon dogs (2–5%) indicated an unspecialized ‘‘villagedog’’ (Shigehara & Onodera, 1984). Amerindian dogsfrom Southwestern sites have cranial Cvs ranging from8 to 17%. This range was thought to represent thepresence of a smaller and a larger breed. The larger dogwas thought to have been used for draught purposes(Haag, 1948). The greatest variation in any populationwas found in remains of Romano-British dogs (Cvrange 11–26%), where a number of small and largedogs breeds is thought to have been present (Harcourt,1974). Historical records suggested that the variety ofdog forms was connected to greater economic affluenceand larger settlement size.Cranial and mandibular Cvs for the kuri are all

under 10%, and the small variability in Facial L.(Cv=6·0%) suggests that it is unlikely there was selec-tion by prehistoric Maori for a short- and long-faceddog, as was suggested by an informant of White’s(1894: 598), who noted that at some Maori settlements‘‘very large heads have been found and at others verysmall ones’’. For the scapula and pelvis, Cvs over 10%can be explained by the early development of theseelements, rather than by genetic or environmentaleffects. The same cannot be said of the limb bones,where there is a consistent pattern, with the highest Cvsfound in the midshaft diameter measurements (Tables3 & 4). It is unlikely that this is due to genetic effects,as Haag (1948) found that morphological variationbetween groups of dogs was present in both cranialand post-cranial elements. Sexual dimorphism is alsoconsidered to be an inadequate cause of midshaftvariation, as the component of variation due to sexin wild canids and modern dogs is between 3 and 7%for most linear dimensions (Hildebrand, 1952; Clark,1995: 211–212).Environmental differences between sections of the

kuri population appear to be the most convincingexplanation, as midshaft dimensions reflect an indi-vidual’s stature and weight, both of which can beinfluenced by nutritional intake (Brothwell, 1981: 163;Reitz & Ruff, 1994: 709). Temporal changes in mid-shaft diameter dimensions were identified betweenearly and late prehistoric sites. However, a detailed


discussion of the causes of this variation is beyond thescope of this paper.Overall, there is no evidence in anatomical traits,

metric attributes or frequency of dental abnormalitiesto suggest that dogs differing in genetic make-up werepresent during New Zealand’s prehistory. It seemsprobable, therefore, either that all kuri were descendedfrom a relatively small founding population, or thatthe skeletal homogeneity was even broader, extendingto central East Polynesian dogs, such as those from theSociety Islands and Marquesas. The latter case wouldprovide supporting evidence for population ‘‘bottle-necking’’ during the west-to-east settlement ofPolynesia. It is important to note that, although the Cvdata does not support differentiation based on mor-phology, prehistoric Maori selection for characteristicssuch as coat colour or behaviour may not be detectablethrough morphometric analysis.

Conclusion

Morphometric analysis of prehistoric Canis familiarisremains is frequently undertaken to provide corollaryarchaeological information about ancient humansocieties (Haag, 1948; Brothwell et al., 1979). Many ofthese studies tend to focus on the cranial elements,leaving the osteology of the post-cranial skeletonlargely unknown (Morey, 1986; Benecke, 1987). Thisapproach limits our understanding of prehistoric dogmorphology and denies the possibility that appendicu-lar skeletal elements can provide useful data for interand intrapopulation comparisons. It is recommendedthat the anatomical and osteometric population par-ameters for as many skeletal elements as possible,cranial and post-cranial, should be examined beforedetailed metric analysis is undertaken. This provides acontext in which the subsequent identification of metricand non-metric skeletal trait variation can be betterunderstood.It is unfortunate that the kuri, as is the case with

other Polynesian domesticates, was not preserved. Incontrast, the free-ranging dingo of Australia and thewild dog of New Guinea, which frequents sub-alpineareas little visited by people, have maintained a largemeasure of their genetic and behavioural identity. Therapid demise of the kuri following the introduction ofEuropean dogs shows that active steps should be takento preserve prehistorically introduced Canis popula-tions before they are irrevocably transformed throughinter-breeding. These animals are often neglected bybiologists and zoologists as they are not indigenousand appear similar to European breeds, a research biaswhich was noted for Polynesia by White as early as1892:

‘‘It is a great pity that travellers in a new country take solittle notice of ordinary or domestic animals, which are thefirst to die out or be modified by interbreeding with their

imported relations; with the result that those who comeafter them addle their brains in a difficult search after relicsof the past.’’ (White, 1892: 554)

In this paper the osteology of cranial and post-cranial elements from the kuri is detailed as a baselinewith which to compare Oceanic assemblages of prehis-toric dogs. Such comparisons will throw light on thepattern of human colonization in the Pacific and clarifythe relationship between Polynesian people and theircommensals.

AcknowledgementsMy thanks to Atholl Anderson, Stuart Bedford andKirsten Lawson for commenting on an earlier draft ofthis paper. I am grateful to the Royal Society of NewZealand for providing funds for field research andto Peter Petchey for photography. The Museum ofNew Zealand authorised the publication of Figure 1(Negative B.3527). Finally, I would like to thank thetwo referees for their helpful and pertinent comments.

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Appendix 1. Archaeological Sites from whichKuri Skeletal Material was Examined

Location Site numberNumber ofmeasurements

Stewart IslandMason Bay D149/? 445Old Neck E48/39 149

South IslandPapatowai G47/50 24Pounawea H47/1 14False Island B44/41 162Akatore I45/2 46Little Papanui J44/1 170Hoopers Inlet I44/13 39Pipikaretu J44/2 51Tarewai Point J44/3 55Kaikais Beach I44/27 473Murdering Beach I44/20 978Long Beach I44/23 176Purakanui I44/21 43Pleasant River J43/1 473Shag Mouth J43/2 1257Redcliffs M36/47 113Moa-bone Point Cave M36/25 54Omihi O32/8 46Peketa O31/15 22Marfells Beach P29/2 118Wairau Bar P28/21 422

North Island

Palliser BayS128/104 &S29/48 451

Paremata R26/122 246Kaupokonui P21/3 505Kohika V15/80 960Tupatika W15/9 160Oruarangi T12/192 127Whangamata T12/2 465Hahei T11/326 19Opito T10/161 284Sarahs Gully T10/167 225Port Jackson S9/53 42Pig Bay R10/22 178Houhora N3/59 703Twilight Beach M2/162 135Tom Bowling Bay na 24

126 G. Clark

Appendix 2. Measurement Description

Cranial Measurements Post-cranial measurements

Skull L. Akrokranion-Prosthion Scapula L. Max. scapula lengthCondylobasal L. Occipital condyles-Prosthion Glenoid L. Glenoid cavity lengthFacial L. Frontal midpoint-Prosthion Glenoid B. Glenoid cavity breadthPalate L. Staphylion-Prosthion Humerus L. Max. humerus lengthNeurocranium L. Akrokranion frontal midpoint Proximal D. Depth of the proximal epiphysisSnout L. Nasion-Prosthion Midshaft (m–l) Midshaft diameter: medio-lateralNasal L. Nasion-Rhinion Midshaft (a–p) Midshaft diameter: anterior-posteriorBizygomatic B. Zygion-Zygion Distal B. Breadth of the distal epiphysisMastoid B. Otion-Otion Radius L. Max. radius lengthPalate B. Max. palate breadth Proximal B. Breadth of the proximal epiphysisBraincase B. Euryon-Euryon Proximal D. Depth of the proximal epiphysisFrontal B. Ectorbitale-Ectorbitale Midshaft (m–l) Midshaft diameter: medio-lateralParaoccipital B. Breadth paraoccipital processes Midshaft (a–p) Midshaft diameter: anterior-posteriorPostorbital B. Breadth postorbital constriction Min. shaft (m–l) Min. corpus diameter: medio-lateralCondyle B. Breadth occipital condyles Distal B. Breadth of the distal epiphysisInterorbital B. Entorbitale-Entorbitale Distal D. Depth of the distal epiphysisMin. Palate B. Measured between P1 and P2 Ulna L. Max. ulna lengthBulla B. Diameter of the auditory bulla Olecranon D. Min. depth across the olecranonSkull H. Skull height with sagittal crest Anconaeus D. Depth across the process aconaeusParietal H. Skull height minus sagittal crest Pelvis L. Max. pelvis lengthAkrokranion H. Basion-Akrokranion Acetabulum B. Breadth of acetabulum: anterior-posteriorOrbital H. Inner height of the orbit Ilium H. Min. height of the ilium body: ventral-dorsalForamen H. Basion-Opisthion Ilium B. Min. breadth of the ilium body: medial-lateralPremolar L. Length of the premolar row Femur L. Max. femur lengthMolar L. Length of the molar row Proximal B. Breadth of the proximal epiphysisP4 B. P4 breadth: cingulum Caput D. Depth of the caput femoris: anterior-posteriorP4 L. P4 length: cingulum Midshaft (m–l) Midshaft diameter: medio-lateralM1 B. M1 breadth: cingulum Midshaft (a–p) Midshaft diameter: anterior-posteriorM1 L. M1 length: cingulum Distal B. Breadth of the distal epiphysisM2 B. M2 breadth: cingulum Tibia L. Max. length of the tibiaM1 L. M2 length: cingulum Proximal B. Breadth of the proximal epiphysis

Midshaft (m–l) Midshaft diameter: medio-lateralMidshaft (a–p) Midshaft diameter: anterior-posterior

Cranic index=Bizygomatic B.#100/Skull L. Distal B. Breadth of the distal epiphysisPalatal index=Palate B.#100/Palate L. Distal D. Depth of the distal epiphysis

Mandibular measurements

Mandible L. 1 Condyle process-Infradentale Corpus P2 H. Ventral corpus-posterior P2 alveoliMandible L. 2 Angular process-Infradentale Premolar L. P1-P4 lengthMandible L. 3 Condylar notch-Infradentale Molar L. M1-M3 lengthM3-P1 L. Cheek tooth row M1 B. M1 breadthM3-P2 L. Cheek tooth row-P1 M1 L. M1 lengthRamus H. Angular process-Coronion M2 B. M2 breadthCorpus M1 H. Ventral corpus-posterior M1 alveoli M2 L. M2 length

osteology of thekurimaori: the prehistoric dog of new zealand

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