COMPARATIVE MORPHOLOGY AND ODONTOMEIRICS OF THE DECIDUOUS OENTITION IN THE RHESUS MONKEY (Macaca muIatta), OLIVE BABOON (Papio anubis) AND KING COLOBUS (CoIobus ponkomos) Thesis for the Degree of M; A. MICHIGAN STATE UNIVERSITY FRANK J. ORLOSKY 1968 IIIIIIIWIIOS L J Umu t)! I HOAGD & suns 800K BINDE'RY EINC. I | ABSTRACT COMPARATIVE MORPHOLOGY AND ODONTOMETRICS OF THE DECIDUOUS DENTITION IN THE RHESUS MONKEY (Macaca mulatta). OLIVE BABOON (Papio anubis) AND KING COLOBUS (Colobus polykomos) by Frank J. Orlosky Odontometric and comparative morphological studies were conducted on the deciduous dentition of the following Old World monkeys: 150 Macaca mulatta; 20 Papio anubis; l7 Colobus polykomos. Tooth measurements were taken with a Boley gauge. A dissection microscope was used in study- ing morphology. Statistical tests were performed with the aid of a computer. Comparative statistical analysis of tooth measure- ments from these animals revealed: the presence of bilateral symmetry in all teeth of the above three genera; the appear- ance of sexual dimorphism in the deciduous molars of all three genera; intraspecific variations of each Species; interspecific differences for Macaca and Papio; statistic- cally significant differences between animals of the three genera considered. Frank J. Orlosky Detailed morphological and comparative observations were presented for all three animal groups. ‘g, mulatta closely resembled g; anubis in many features. whereas both groups differed from the morphology of §;_polykomos. Dis- tinguishing features for each group were summarized and diagnostic criteria were suggested. COMPARATIVE MORPHOLOGY AND ODONTOMETRICS OF THE DECIDUOUS DENTITION IN THE RHESUS MONKEY (Macaca mulatta). OLIVE BABOON QPapio anubis) AND KING COLOBUS (Colobus polykomos) BY Frank J. Orlosky A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF ARTS Department of Anthropology 1968 ACKNOWLEDGEMENTS I would like to express my appreciation and indebted- ness to Dr. Daris R. Swindler. the chairman of my graduate committee. Dr. Swindler supplied the primate skeletal and cast material used in this thesis as well as the guidance and encouragement without which this work could not have been completed. I want to thank Dr. James Brown. my second com- mittee member. for his kind advice during the writing of this thesis and especially for aid with the statistical analysis of data. I wish to express appreciation to Charles Wood for his patient assistance in the preparation of the figures and to Roberta English for secretarial aid. I am grateful to Dr. James Gavan. the former director of the Fort Johnson Primate Colony. for allowing me to study the Macaca mulatta cast ma- terial which was processed under his supervision. I wish to thank Susan Frost for her encouragement and assistance during the final stages of this work. ii TABLE OF CONTENTS Page ACKNOWLEDGEMENTS . . . . . . . . . . . . . . . . . . ii LIST OF TABLES . . . . . . . . . . . . . . . . . . . vi LIST OF FIGURES. . . . . . . . . . . . . . . . . . . vii INTRODUCTION . . . . . . . . . . . . . . . . . . . . 1 REVIEW OF LITERATURE . . . . . . . . . . . . . . . . 4 MATERIALS AND METHODS. . . . . . . . . . . . . . . . 7 Definitions and Methods of Measurements. . . . . 8 Definitions and Methods Regarding Morphology . . lO ODONTOMETRICS OF THE DECIDUOUS TEETH OF MACACA MULATTA. PAPIO ANUBIS. AND COLOBUS POLYKOMOS . . 12 Correlation of Right to Left Sides of Dental Arch . . . . . . . . . . . . . . . . . . . . 13 Sexual Dimorphism. . . . . . . . . . . . . . . . lS Macaca mulatta . . . . . . . . . . . . . . . lS Papio anubis . . . . . . . . . . . . . . . . l6 Colobus polykomos. . . . . . . . . . . . . . 18 IntraSpecific Variation. . . . . . . . . . . . . l9 InterSpecific Differences. . . . . . . . . . . . 21 Intergroup Differences . . . . . . . . . . . . . 23 Relative Sizes of Teeth. . . . . . . . . . . . . 23 iii Table of Contents (continued) DESCRIPTIVE MORPHOLOGY OF THE DECIDUOUS DENTITION OF MACACA MULATTA. PAPIO ANUBIS. AND COLOBUS POLYKOMOS. . . . . . . . . . The Maxillary Dentition. . . . . The The The The The The Upper Central Incisor. . Macaca mulatta . . . . . Comparative Observations Upper Lateral Incisor. Macaca mulatta . . . . Comparative Observations Upper Canine . . . . . . Macaca mulatta . . . . . Comparative Observations Upper First Molar. . . . Macaca mulatta . . . . Comparative Observations Upper Second Molar . . . Macaca mulatta . . . . . Comparative Observations Mandibular Dentition . . . . The The The Lower Central Incisor. Macaca mulatta . . . Comparative Observations Lower Lateral Incisor. Macaca mulatta . . . Comparative Observations Lower Canine . . . . Macaca mulatta . . . Comparative Observations iv Page 27 28 28 28 31 32 32 34 36 36 37 39 39 43 46 46 50 53 53 53 54 55 55 57 58 58 6O Table of Contents (continued) Page The Lower First Molar. . . . . . . . . . . . . 6l Macaca mulatta . . . . . . . . . . . . . . 61 Comparative Observations . . . . . . . . . 65 The Lower Second Molar . . . . . . . . . . . . 66 Macaca mulatta . . . . . . . . . . . . . . 66 Comparative Observations . . . . . . . . . 69 DISCUSSION . . . . . . . . . . . . o . . . . . . . . . 72 SUMMARY. . . . . . . . . . . . . . . . . . . . . . . . 84 APPENDIX . . . . . . . . . . . . . . . . . . . . . . . 85 LITERATURE CITED . . . . . . . . . . . . . . . . . . . 103 LIST OF TABLES Table 1. Correlation of right and left side dentitions within each species. . . . . . . . . . . . . 2. Comparison of male and female tooth measure- ments within each species. . . . . . . . . 3. Comparison of tooth measurements of Macaca mulatta. Papio anubis. and Colobus polykomos 4. Relative sizes of teeth within each Species. . Page 86 89 95 98 LIST OF FIGURES Figure Page 1. Comparison of mesio-distal tooth dimensions in Papio anubis and Papio ursinus at the t 0.05 confidence level . . . . . . . . . . . . . . . 99 2. Comparison of bucco-lingual tooth dimensions in Papio anubis and Papio ursinus at the i 0.05 confidence level . . . . . . . . . . . . . . . lOO 3. Morphology of the maxillary deciduous dentition. 101 4. Morphology of the mandibular deciduous dentition 102 vii COMPARATIVE MORPHOLOGY AND ODONTOMETRICS OF THE DECIDUOUS DENTITION IN THE RHESUS MONKEY (Macaca mulatta). OLIVE BABOON (Papio anubis) AND KING COLOBUS (Colobus polykomos) INTRODUCTION The study of fossilized teeth has played a major role in the advancement of knowledge concerning the evolu- tion of Hominids as well as of the nonhuman primates. This is true simply because the primate paleontological record consists largely of fossilized teeth. Every fossil specimen. therefore. receives close study and repeated descriptions. Studies of the dentition of living primates also play an integral role in understanding the evolution of man. It is through such studies that the morphological and metric var— iability existing in the dentition of living groups of pri- mates may be discerned. Having established the parameters of variability for each primate species. characteristic dif- ferences between taxonomic groups of primates may be realized and the positioning of fossil specimens may be more thoroughly understood. Realizing the importance of dental studies of the living nonhuman primates. it is startling to discover that little has been written concerning the teeth of these animals. This surprising state of affairs has been noted by Ashton and Zuckerman (1950b; 213). Straus (1955; 130). Butler (1963; 2). Frisch (l963; l6). Swindler (1963; 104). Biggerstaff (1966; 23). and others. The above authors wrote in reference to the permanent dentition; the situation is far worse in regard to the deciduous dentition. Very little has been written con- cerning the deciduous teeth of any nonhuman primate; the studies which have been published to date will be reviewed in the following section. The purpose of this thesis is to present data on the morphology and odontometrics of the Macaca mulatta deciduous dentition and to compare these findings with the existing studies of the deciduous teeth of other Cercopithecidae. The results will also be compared with observations and measure- ments taken from casts of one other genus of Cercopithecinae (Papio anubis) and one genus of Colobinae (Colobus polykomos). This thesis is largely an attempt to discern the de- gree of variability present in the deciduous dentition of each of the above groups of animals and to establish diagnostic differences between species of two genera and be- tween members of different genera. The present study is necessarily limited to the above three genera and the scarc— ity of previous data reduces the possibilities for valid generalizations. The results of this study. however. may be of future value to the paleontologist. as well as to the physical anthropologist. for interpreting fossil material and in differentiating between existing groups of Cercopithe- cidae. REVIEW OF LITERATURE A review of the literature revealed several reports dealing in part with the deciduous dentition of Old World monkeys. It was found that early dental studies were con- cerned largely with the general form. identities and homol- ogies of the deciduous teeth. This is the nature of the general remarks of Owen (1840-1845; 441). Flower (1867-1869; 270). and Tomes (1894; 471). The remarks of Bennejeant (1953; 85) and James (1960; 168) also fall into this cate- gory. Bennejeant merely stated that the temporary molars of Cercopithecids are bilophodont while James mentioned. as had Tomes. the absence of sexual dimorphism in the size of the deciduous canine. Several studies have been concerned with the odonto- genesis and eruption of the deciduous teeth of Old World monkeys. The studies of Swindler and McCoy (1964; 1243) and Swindler et a1. (1968; 167) were concerned with molar calci- fication in M; mulatta and Papio anubis respectively. Schultz (1933; 20). Marshall (1933; 85). and Hurme and van Wagenen (1953; 291) were concerned with the eruption of the deciduous 4 teeth of M; mulatta. Likewise. Nissen and Riesen (1945; 265) concentrated on the eruption of the deciduous teeth of the chimpanzee. The morphology of the deciduous dentition of Old World monkeys has been considered by several authors. Remane (1951; 162) considered the lophodont variability of EEEEQ' pithecus and Colobus. Okerse (1959; 222) presented a typo- logical description of the deciduous dentition of the Vervet Monkey (Cercopithecus aethiops). Freedman (1957; 132) briefly discussed the general morphology of the deciduous teeth of Papio ursinus. Several studies related to the metric variability of deciduous teeth have been conducted. Auskaps and Shaw (1957; 432). in their work on M; gynomolgus. included measurements of three juveniles and eleven animals of mixed dentition; they concluded that there were no sexual differences in the size of teeth. Freedman (1962; 229) presented data on the metric variability of the deciduous teeth of g; ursinus and concluded there was no difference between the sexes in the size of the milk teeth. Ashton and Zuckerman (1950b; 212) included measurements of the milk teeth of Cercopithecus aethiops sabaeus° Biggerstaff (1966; 231) compared the deciduous molar measurements of M; mulatta and_ML speciosa and found that the two Species could be differentiated sta- tistically° Ashton (1956; 121) studied the sexual differ— ences of the milk teeth of the chimpanzee and gorilla and found an absence of dimorphism in the former but a presence of sexual differences in the latter. At the present time. the literature lacks detailed morphological. metric. or comparative studies of the decid— uous dentition of any Old World monkey. The following work was undertaken in an attempt to partially fill this void. MATERIALS AND METHODS The deciduous dentition of Macaca mulatta was studied from skulls collected by D. R. Swindler. The sample consisted of 150 animals. 75 males and 75 females. the sex of whom was determined at the time of decapitation. No animal was in- cluded in this study if the sex was not known. .M; mulatta range from the west coast of Bombay and northward from the valley of the Godaveri to the Himalayas (Pocock 1910; 230). Specific data as to the exact origin of the macaque skulls is lacking. In addition to the skull collection. 90 dental casts taken from a'M; mulatta colony located at Fort Johnson. South Carolina. were studied for detailed morphological var- iability. In each case. jeltrate impressions were made and albastone was used as the casting material. It has been demonstrated by Swindler et a1. (1963; 105) that no appre- ciable error occurred in measurements made from such casts. The Fort Johnson casts were taken serially from 30 animals (16 females; 14 males) as the deciduous teeth were erupting. 'Dhus. the dentition of these animals were almost totally unworn and presented the best possible material for detailed morphological observations. The deciduous dentition of Papio anubis was studied from dental casts of 20 animals (12 females; 8 males) cap- tured 120 miles east by southeast of Nairobi. Kenya (Swindler 1967; 134). The deciduous teeth of Colobus polykomos were studied from casts taken of specimens in the collections of the Smith- sonian Institute and the Chicago Field Museum. Tappen (1960; 107) reported on the wide range of g; polykomos which extends from Angola through the Congo forest and includes the forest outliers in the savanna region adjoining the Congo. The ma- jority of casts used in this study were taken of Specimens originating from the countries of Ethiopia. Sudan. and Kenya. Two specimens originated from the Congo and one from Ghana. The total sample consisted of 17 animals (10 males; 7 females). Definitions and Methods of Measurement Measurements were taken with a Boley gauge equipped with a Vernier scale which permitted readings within the nearest 0.1 mm. Repeated measurements on the same teeth. six months after the original readings. revealed an average difference in measurements of only 0.13 mm. Mesio-distal and bucco-lingual (labio—lingual)mea- surements were taken of each tooth in every Specimen. Both of these measurements will Simply be referred to as diameters to avoid confusion which has arisen in the use of terms such as breadth or thickness (Selmer-Olsen 1949; 23). The mesio—distal diameter is defined as the greatest distance between mesial and distal contact points on the oc- clusal surface. Where no mesial or distal contact points are present. the greatest measurement along the mesio-distal axis of the tooth was taken. Both the upper and lower lateral in- cisors. however. are positioned obliquely in their alveoli. To avoid disproportionately large readings. the mesiobdistal measurements of these teeth were taken vertical to the long axis of the tooth rather than perpendicular to the occlusal surface (Selmer-Olsen 1949; 24 and Wheeler 1956; 25). The bucco—lingual (labio-lingual) diameter is defined as the greatest bucco-lingual measurement taken perpendicular to the mesio—distal diameter. This measurement does not lie in the same horizontal plane as the mesio-distal measurement. 10 Definitions and Methods Regarding Morphology A dissection microscope (powers from .7X to 3X) was used in studying morphology. Standard terminology was used to describe the deciduous dentition. The following defini- tions are included as an aid to the reader. Buccal surface: Labial surface: Lingual surface: Mesial surface: Distal surface: Tubercle: Ridge: Marginal ridge: pertaining to the surface of a molar which is next to the cheek pertaining to the surface of an anterior tooth which is toward the lips pertaining to the surface of a tooth toward the tongue that surface of a tooth which is toward the median line of the dental arch that surface of a tooth which is away from the medial line of the dental arch a small elevation on the crown of a tooth due to extra enamel any elongated elevation or crest on the surface of a tooth a ridge which forms either the mesial or distal margins of the occlusal surface of a molar; also a ridge which forms the mesial or distal margins of the lingual surface of an anterior tooth Developmental groove: a groove located between cusps which indicates the coalescence of the lobes Central developmental groove: that groove which passes mesio-distally between the buccal and lingual cusps Buccal and lingual developmental grooves: developmental grooves which pass between the buccal cusps and lingual cusps reSpectively ll Fossa: a rounded depression or concavity on the surface of a tooth Mammelon: one of three small rounded protuberances on the incisal edge of the incisors In addition to the basic definitions above. the Cope- Osborn cusp designation was employed throughout this work. The following lists present the COpe—Osborn designation and the position of each cusp on the occlusal surface for both the maxillary and mandibular molars. Maxillary Molars Mandibular Molars paracone : mesio—buccal protoconid : mesio-buccal protocone : mesio-lingual metaconid : mesio-lingual metacone : disto—buccal hypoconid : disto-buccal hypocone : disto-lingual entoconid : disto—lingual ODONTOMETRICS OF THE DECIDUOUS TEETH OF MACACA MULATTA. PAPIO ANUBIS AND COLOBUS POLYKOMOS G. G. Simpson (1943; 151) has elaborated on the im— portance of quantitative data and of statistical analysis to taxonomy. Simpson maintains that populations of animals. rather than types. are of central concern in defining taxa- nomic categories; especially in the definition of species. In regard to the study of animal p0pu1ations. quantitative. or quantifiable. characteristics are very useful since such observations are more accurately repeatable. relatively pre— cise and easily subjected to statistical analysis. The sta- tistical analysis of dental measurements is therefore a con- venient and valuable indicator of the limits of dental varia- bility of animal groups and of diagnostic differences between various groups. The major purposes of this section are to determine the degree of metric variability within each group of Old World monkey under consideration and to determine the metric boundaries separating these groups. 12 13 Measurements of the deciduous dentition of the three groups of Old World monkeys were taken as described in the previous section. These data were then transferred to For- tran sheets. punched on cards and submitted to a computer. The card punching. varifying. and programing were performed by the computer laboratory service at Michigan State Univer- Sity. Correlation of Right to Left Sides of Dental Arch The existence of a high degree of correlation be— tween right and left sides of primate dentitions has been demonstrated for the permanent teeth of several animals by Ashton and Zuckerman (1950; 474) and by Schuman and Brace (1955; 63). The symmetry of the deciduous dentition of the animals presently under consideration was checked by taking measurements of the teeth of both sides of the dental arch and calculating the sample correlation coefficient (r) by means of a computer. Table 1 presents the computed sample Size. mean and simple correlation value (r) for each tooth of each animal species. The simple correlation values for each of the right-left combinations was high for all three 14 groups of animals. The null hypothesis (Ho:p=0. HA:p#0). or the hypothesis of no correlation between right and left sides. was easily tested for each combination by referring to Table 7.6.1 of Statistical Methods (Snedecor 1956; 174) under the appropriate degrees of freedom. In every case in Table l. the null hypothesis was rejected at the .05 level. In addi- tion. all values of r were significant at the .01 level ex- cept the (M-D) il measurements of Papio anubis and Colobus polykomos. The teeth of the right side. therefore. did not Sig- nificantly differ from the left side in either measurement. The common practice of using only one side of the dental arch for comparisons or for testing hypotheses will therefore be justifiably used in subsequent portions of this section. The sample sizes (N) presented in later tables repre- sent only the right side of the dental arch. As pointed out by Ashton and Zuckerman (1950; 474). it is not justifiable to treat right and left tooth measurements as independent varieties. i.e. the two sides should not be added to obtain a larger N. 15 Sexual Dimorphism The absence of sexual dimorphism in the deciduous dentition of nonhuman primates has been reported by Ashton (1956; 121). Auskaps and Shaw (1957; 433). James (1960; 168). Freedman (1962; 232) and it has been assumed by Biggerstaff (1966; 232) and others. To check for sexual dimorphism in the deciduous dentition of the animals presently under con— sideration. the null hypothesis (Ho:ul=u2. Hl : ul # u2). or the hypothesis of no difference between the sexes. was tested by performing t tests on the measurements of each tooth of each Species. The sample size. mean. standard deviation. coefficient of variation. standard error and t value for each male-female combination are presented in Table 2. Macaca mulatta The null hypothesis was not rejected for the mesio— distal and bucco-lingual measurements of the ML_mulatta an- terior deciduous teeth. Thus. the measurements of the an- terior teeth did not differ Significantly in size between males and females. The null hypothesis was rejected. however. for both diameters of the maxillary and mandibular molars. 16 i.e. the t values were significant at the .05 level. All but one of the t values were significant at (.001. The males were. in all instances. larger than the females. The differ- ences between the mean values were as great as .19 mm which is significant Since the sample size was relatively large. Sexual dimorphism therefore exists in the mesio- distal and bucco-lingual dimensions of the deciduous molars 0f.fl; mulatta. This should be accounted for when comparing this animal's dentition with other groups. i.e. the absence of sexual dimorphism should not be assumed 3 priori. This is especially true if discriminative tests are performed on comparative data which are sensitive to such statistically significant differences. Papio anubis As in M; mulatta. the null hypothesis was not rejected for the anterior teeth of P; anubis. i.e.. significant differ- ences did not appear between the sexes in either measurement. The incisors of the male animals. however. were represented by only one Specimen. The t values computed do not. there- fore. constitute accurate estimates of the real differences between males and females in these measurements. The female 17 sample is relatively large and. for this reason. these data are included in the table. The null hypothesis was not rejected for the first deciduous molars (at the .05 level). but was rejected for the bucco-lingual dimension of both the upper and lower second molar. The null hypothesis was also rejected for the mesio-distal measurement of the upper second molar. The differences which appear between the means of males and females range from .509 mm to .707 mm in the above measurements of the second molars. The males. again. were larger than the females in both dimensions. Freedman (1962; 232) tested for sexual dimorphism in the lengths of the deciduous molars in}:é ursinus. Of the t tests performed on the four mean measurements. two were significant at the .05 confidence level. while two were not significant. Freedman concluded. however. that sexual di- morphism was not present in the size of deciduous teeth of P; ursinus. Freedman's data on P;_ursinus showed that the greatest difference in mean values occurred in the mesio—distal measure- ment of the upper second molar. This undoubtedly was one of the two t values which proved significant. It is interesting 18 to note that this was the only mesio-distal measurement which was significant in the P; anubis sample. Freedman did not report of tests performed on bucco-lingual measurements. The present study. at least in part. agrees with Freedman's find- ings on_P; ursinus. It is suggested. however. that some de- gree of sexual dimorphism does exist in the deciduous second molars of both Species of Papio. Colobus polykomos AS with g; anubis. the null hypothesis was not re- jected for the deciduous anterior teeth and the first de— ciduous molar. i.e. the measurements of these teeth did not demonstrate sexual dimorphism. The upper second deciduous molar likewise did not demonstrate a significant difference between sexes in either measurement. The t values of both measurements of the second mandibular molar were significant at the .05 level. i.e. there was a significant difference between males and females in both dimensions and the null hypothesis was rejected. Unlike the above two groups. how- ever. the females were larger than the males. 19 Intraspecific Variation The magnitude of variation present in a quantitative trait within a population may be analyzed in several ways. The least complex and most direct. in the author's opinion. is by studying coefficient of variation values. The coeffi— cient of variation is Simply the relative standard deviation which is calculated by dividing the standard deviation by the mean (Snedecor 1956; 44). The resulting values 'correct' for the larger numerical variances usually associated with larger teeth and provide a measure of relative variance which can be directly compared. The degree of intraspecific variation of each tooth dimension for each sex of each Species is presented in the form of coefficients of variation (C.V.%) in Table 2. Exam- ination of these values revealed the following variatiOn trends: 1) In general. the dimensions of the molars of each Species were less variable than the anterior teeth. This might be expected Since these teeth are more subject to wear which might affect the measurements taken. This is probably only a contributing factor. however. Since the incisors were relatively less variable in the bucco—lingual dimension which is affected more by attrition. 2) Within each species. the maxillary and mandibular first molars tended to be more variable than the 20 upper and lower second molars. This trend existed in both mesio-distal and bucco-lingual dimensions and in both sexes. A few exceptions were present in the data. most notably in the mesio-distal dimen— sion of the molars of M; mulatta. In each instance. however. the difference between the C.V° values was less than .5%. 3) Within each Species. the lateral incisors tended to be more variable than the central incisors. Gener- ally. there were relatively large differences between the C.V. values for these two teeth. Of the 24 com- parisons in Table 2. there were four exceptions to this trend. The above trends partially support Butler's dental 'field' concept (Butler 1939; 1). The central incisors tended to be more stable while the lateral incisors showed greater degrees of variation. If the first deciduous molars are con- sidered to be the center of the 'deciduous molar field.I the data does not support the hypothesized mesial to distal in- crease in the degree of variability. If. however. the de- ciduous molars are considered as a unit with the permanent molars. the center of the 'molar field' would be on the first permanent molars and the degree of variability would be ex- pected to increase from distal to mesial. 21 Interspecific Differences The amount of Odontometric data published on the de- ciduous dentition of the Old World monkeys is very small. Of the present publications. the only apprOpriate works for making interspecific observations are Freedman's study of g; ursinus (1962; 229) and the comparative work of Bigger- staff (1966; 231) on M; mulatta and M; speciosa. Biggerstaff's work was of limited comparative value. First. only the four deciduous molars of the two species of macaque were considered. Secondly. the sexes of the animals were unknown and the Specimens were therefore lumped together. In the present study it was shown that dimorphism was present in just these two teeth. Thirdly. the measurements of the right and left sides were added together producing N values greater than the actual sample Size. Nevertheless. Bigger- staff's data on the molars of M; mulatta were quite similar to the values obtained in the present study. The maximum difference between the mean values of Biggerstaff's sample and either sex of the present study was .3 mm. In the present study. however. the magnitude of variability (at the i .05 confidence level) was less and variations existed be- tween the sexes. The data of the present study supports 22 Biggerstaff's conclusions concerning the differences between the molar dimensions of the two macaque species. Papio anubis and Papio ursinus are compared at the : .05 confidence level in Figures 1 and 2. In the present study. the specimens of both sexes were combined because the absence of sexual dimorphism (except in three measurements of the two second molars) has been demonstrated and because the male sample was small. especially for the incisors. The combination of data from both sexes tends to numerically in- crease the mean and range values of the female Specimens. A more conservative assessment of the differences between the two species should therefore be obtained if the combined sex data is compared with the female data of Freedman's work. Examination of Figures 1 and 2 revealed that the confidence intervals for the bucco-lingual dimensions of all teeth of both species. overlapped. Distributional over- lap was not present. however. in the mesio-distal dimension of six teeth. The distance between the confidence intervals was greatest for the upper and lower second molars. Of the deciduous teeth of Papio. the second molars appear to be the most useful for metrically differentiating between species. This agrees “nth Biggerstaff's conclusions on the macaques. 23 Intergroup Differences To determine the presence or absence of significant differences between the tooth measurements of the groups of animals used in this study. the hypothesis of equal means ( HO: 1 = “2 = u3. Hl : ul # u2 = u3 ) was tested by using an F test. Table 3 presents the F statistics and the approx- imate significance level for the right upper and lower denti- tions of the three animal groups under consideration. In each case. the hypothesis of equal means was rejected. Thus. the results showed that the deciduous dentition of the three groups of animals differed greatly in size; the teeth of Papio anubis were uniformly larger than those of Macaca mu- latta which were in turn correspondingly larger than the teeth of Colobus polykomos. The probability of the F statis- tic for each tooth was (.0005 for both measurements. Thus. there was a very significant difference in tooth dimensions between the above three groups of animals. Relative Sizes of Teeth The absolute differences in Size between various species and genera are important for taxanomic and evolu— tionary considerations. The relative sizes of a species 24 dentition and differences in trends between Species and genera are also important for similar reasons. The relative sizes of the teeth of the three species under consideration can be determined by studying Table 2. These data have been summarized in Table 4. To this has been added Freedman’s data on Papio ursinus (1962; 229). The dif- ferences between each successive tooth dimension is presented in parenthesis below each inequality sign. The Papio anubis male tooth dimension sequence was based on a small sample; the female sequence is therefore assumed to be more repre- sentative of the species. There were no sexual differences in the mesio-distal dimension sequences. although Slight variations existed in the differences between means. The sequences of the mesio- distal dimensions of the mandibular dentitions were the same for all four species presented in Table 4. Macaca mulatta and Papio anubis demonstrated the same maxillary mesio—distal dimension sequence. ‘2; ursinus. however. appeared to differ from the above two Species in that it had a lateral maxillary incisor which was larger than the upper canine. The differ- ences between means among females. however. was smaller than the error of measurement. The differences between males was 25 somewhat larger. but the sample size was only half that of the females. It is probable. therefore. that this does not constitute a valid interspecific difference. Colobus pg_y— komos demonstrated a maxillary canine which was larger than either of the maxillary incisors. The differences between successive teeth were large and this probably constitutes a Species trend and a valid difference between this species and the above species of Cerc0pithecinae. The bucco—lingual dimension sequence was similar for the three Cercopithecinae for which data are presented. The sequence for M; mulatta was: mZ-m —il-(cl.i2); while for the two species of Papio the canine was found to be larger than the upper second incisor. In both cases. how— ever. the numerical difference between the mean values of the canine and incisor was very small. The sequence for the bucco-lingual tooth measurements of g; polykomos was: mZ—ml—cl-il-iz. The differences between successive teeth were large and this may also constitute a valid differenti- ating trait of this species. The bucco—lingual mandibular sequence for M; mulatta Showed that the lower second incisor was smaller than the central incisor and the canine; the latter two. however. 26 demonstrated a very small difference between mean values. In the sequence for the mandibular teeth of P; anubis the canine was smaller than either of the incisors. which dif- fered only slightly in their bucco—lingual dimensions. This relative size difference may constitute a significant dif- ference between these two groups of animals. ‘E; ursinus demonstrated a slight sexual difference in sequences of mandibular bucco-lingual tooth measurements. In males. the central incisor was Slightly smaller than the canine or lateral incisor; while in the females the central incisor was larger than both the canine and lateral incisor. In either case. the sequence did not correspond to that of _§; anubis. Since the canine was not smaller than the incisors. This trend may be of value in differentiating between these two Species of Papio. The mandibular bucco—lingual sequence of g;_pglykomos was different from all three of the above animals. In both sexes. the canine was larger than both of the incisors. In each case. there was a relatively large difference between mean values of successive teeth. In addition to the above. ‘9; polykomos was further differentiated from M; mulatta by the greater size of the lateral incisor relative to the medial incisor. DESCRIPTIVE MORPHOLOGY OF THE DECIDUOUS DENTITION OF MACACA MULATTA. PAPIO ANUBIS AND COLOBUS POLYKOMOS No detailed study of non-metric morphological char- acteristics of the deciduous teeth of any Old World monkey was found in the existing literature. The observations recorded to date were briefly reviewed in a previous sec- tion. None of these. however. offer sufficient details to distinguish between various groups of Old World monkeys. The following section is concerned with the dental. nonmetric. morphological similarities and differences of three groups of Cercopithecidae: Macaca mulatta. Papio anubis. and Colobus polykomos. The sample of g; anubis upon which comparative observations were made consisted of 12 females and 8 males. while the sample of Q; polykomos was composed of 10 males and 7 females. Because of wear. however. a particular observation could not be made on all of the above casts. The observations of the M; mulatta skull collection were supplemented by study of dental casts taken of the Fort Johnson Rhesus colony. 27 28 The Maxillary Dentition THE UPPER CENTRAL INCISOR Macaca mulatta The upper central incisor presented four surfaces: labial. lingual. mesial. and distal. The mesial and distal surfaces were triangular in outline with the apex at the in- cisal edge; cervically. these surfaces sloped disal and me- sial respectively (Fig. 3). Both surfaces were smooth and rounded in both directions. The cervical line of both as— pects curved incisally at a sharp angle. The labial surface was roughly trapezoid in outline with the base at the incisal edge. The labial surface was also smooth and it was rounded both mesially and distally. Detailed observations of the lingual surface were made of the Fort Johnson cast collection. The lingual sur- face was triangular in shape with the apex at the cervix of the tooth. Upon eruption. there was no evidence of mammelons (on the incisal edge; these have been reported during calci- fication (Swindler. 1964; 1243). The lingual surface was bordered by mesial and distal marginal ridges which were continuous with an incisal ridge (Fig. 3. A). The incisal 29 ridge sloped both mesially and distally from a high point located approximately 1/3 the distance from the disto—incisal angle. The mesial and distal marginal ridges traveled cerv- ically. sloped distally and mesially respectively. and term- inated as they contacted a diagonal lingual ridge. The lin— gual ridge had a variable expression which resulted in three varieties of lingual morphology. These ranged from a smooth surface to a large lingual tubercle. Three categories of variation were discernible: 1) Smooth—-very slight expression of ridge giving the appearance of a smooth lingual surface. 2) Ridge-—a well developed ridge traveling from the mid- point of the cingulum to a point cervical to the incisal edge and a variable distance from the mesial-incisal angle. 3) Tubercle——a broad. thick elevation that protruded more lingually than the diagonal ridge. It was positioned along the midline of the tooth and generally traveled vertically from the cervix to approximately 1/2 the vertical height of the tooth. The percentage of occurrence is charted below for each type of morphology for each sex in the Fort Johnson cast collection. Since the expression was symmetrical. only one side of the dental arch is recorded. 3O Variations of the Lingual Surface of the Upper Central Incisors of Macaca Mulatta Sex Smooth Ridge Tubercle M 2 10 1 (15.4%) (76.9%) (7.7%) F 2 12 2 ‘ (12.5%) (75.0%) (12.5%) As shown here. the ridge morphology was the most frequently occurring type in both sexes. The smooth and tubercle varieties occurred equally infrequently in both sexes. In addition to the Fort Johnson cast collection. the cervical portion of the lingual surface was observed in 76 M£_mulatta skulls. Although the cervical portion of a diagonal ridge was observed in all 76 skulls. the shape and termination of the ridge on the incisal third of the lingual surface could not be determined because of wear. (One fe- male skull showed a fusion of the two upper incisors. The fusion was more completed on the left side and. on both Sides. included fusion of the roots.) 31 Comparative Observations In general. the morphology of the upper central in- cisor of E; anubis resembled that of M; mulatta. Aside from metric differences. however. there were several morphological features which readily differentiated the two groups of ani- mals. The labial and lingual surfaces of the E; anubis upper central incisor were more rhomboid in general outline since the cervical portion was relatively larger than that of M; mulatta (Fig. 3. b). The lingual morphology was basically similar in both animals. As in M; mulatta. mesial and distal ridges bordered the lingual surface and dissipated as they approached a lingual diagonal ridge. The diagonal ridge was positioned as in the Rhesus monkey. but it appeared not to continue as far incisally (Fig. 3. b). Because of wear. how— ever. these observations could be made on only seven speci- mens and the terminations of the diagonal lingual ridge could not be absolutely determined. The upper central incisor of g; polykomos was consid- erably different in morpholOgy from both M; mulatta and}:é anubis. The labial and lingual surfaces more closely re- sembled the rhomboid general outline of the baboon upper central incisor. The most diagnostic characteristics. 32 however. occurred in the details of the lingual surface. The marginal ridges of the lingual surface continued uninterrupted along the cervical portion of the tooth and formed a cervical ledge which was undercut labially to a variable degree. A small tubercle often appeared approximately midway along this cervical ledge. There was no diagonal ridge present on the lingual surface. The lingual surface was divided approxi— mately in half by a vertical. small elevation which created a slight depression on both of its sides. This elevation ex— tended for a variable distance cervically and often created slight depressions on the incisal edge. These depressions probably represented remnants of mammelons (Fig. 3. c). De- tailed observations were possible on 10 specimens: 7 males. 3 females. THE UPPER LATERAL INCISOR Macaca mulatta The upper lateral incisor was positioned obliquely in the posterior half of the premaxilla bone. The root traveled dorsal and cranial. and crossed the maxilla— premaxilla suture so that the superior half of the root 33 rested in the maxilla. The root was flattened labio- lingually and had a lingual curve at the apical end. The tooth presented four surfaces: labial. lingual. mesial. and distal. The distal surface was triangular with the apex at the incisal edge. and it was curved mesially in both directions. The distance from the cervix to the incisal edge was small because of the continuation of the incisal edge disto—cervically and because of the angulation of the tooth. The mesial surface was triangular in shape and was slanted mesially (Fig. 3. d). The surface was smooth and triangular; it curved lingually in both directions. The in- cisal edge sloped superiorly in both directions from a point approximately 1/3 from the: mesio-incisal angle. The mesial slope was gradual; Unadistal lepe was more curved and term- inated as the incisal edge met with the distal surface (Fig. 3). Because of the oblique position of the upper lateral incisor and because of the mode of occlusion. the mesio— incisal portion of the lingual surface was greatly effected by wear. The details of the lingual surface were therefore largely obliterated in the Macaque Skull collection. The 34 following statements are therefore based largely on obser- vations of the less worn Specimens of the Fort Johnson cast collection. The lingual surface was roughly triangular in outline with the base on the incisal ridge (Fig. 3. d). Me- sial and distal marginal ridges were present along the re— spective borders of the lingual surface; these ridges curved slightly distally and mesially respectively to end at the cervix of the tooth. A diagonal lingual ridge traveled mesio-incisally from the cervical termination of the two marginal ridges and terminated at a point cervical to the incisal ridge and a variable distance from the mesio—incisal angle (Fig. 3. d). This ridge occasionally traveled more mesially so that it almost joined with the mesial marginal ridge and thus gave the appearance of a smooth lingual sur- face. The Shape and position of the above ridge was similar to that of the upper central incisor. No lingual tubercles and no differences in morphology between the sexes were ob- served. Comparative Observations The morphology of the upper lateral incisor of P; anubis agreed well with the above description of the maxillary 35 second incisor of M; mulatta. As in M; mulatta. the labial and lingual surfaces were triangular in shape with the apex at the cervix of the tooth. Likewise. mesial and distal marginal ridges were present on the lingual surface. These terminated as they approached the diagonal lingual ridge which traveled from the cingulum to a variable point cranial to the incisal edge (Fig. 3. e). Because of wear. the lin- gual morphology could be observed in only six Specimens. In these six. however. the diagonal ridge traveled only a slight distance incisally. This was similar to the extent of the lingual ridge of the upper central incisor of B; anubis. In g; polykomos. the lingual and labial surfaces were triangular in outline but the apex was located at the occlusal surface of the tooth. The lingual surface was bordered by mesial and distal marginal ridges which were continuous with the incisal ridge. The marginal ridges. as in the upper central incisor. continued uninterrupted along the cervical portion of the lingual surface. Unlike g; anubis and M; mulatta there was no diagonal lingual ridge present on the lingual surface (Fig. 3. f). The lingual surface was smooth and convex; also. there were no vertical elevations on this surface as in the upper central incisor of this animal. 36 THE UPPER CANINE Macaca mulatta The upper canine presented four surfaces: labial. lingual. mesial. and distal. All four surfaces were roughly triangular in shape with the base at the cervix of the tooth. The mesial surface was smooth and rounded both labially and lingually; it was continuous with the labial surface. The incisal edge sloped both mesially and distally; the mesial Slope was steep and it was continuous with the mesial sur— face. while the distal slope of the incisal edge was more gradual and terminated at the distal surface. As the distal portion of the incisal edge approached the distal surface it traveled horizontally for a short distance. and thus formed a slight distal ledge or tubercle. On the labial surface there was a slight depression mesial to this distal bulge. Aside from this depression. the labial surface was smooth and rounded in both directions. The distal surface consisted of a small triangular section of enamel cervical to the distal termination of the incisal edge. The incisal portion of this surface blended into the distal portion of the incisal edge. The cervical portion was smooth and rounded mesially in both directions. 37 The lingual surface was bordered by mesial and dis- tal ridges. These ridges passed cervically from the tip of the cusp. traveled distally and mesially respectively and terminated as they approached a vertical lingual ridge. The lingual ridge bowed mesially from approximately the midpoint of the cervix of the tooth to the apex of the crown and di- vided the lingual surface into two unequal portions (Fig. 3. 9). Aside from slight variations in the slope of the lingual ridge and in the degree of expression of the distal ledge. there were no morpholOgical variabilities in the above pattern. No sexual differences in the above morpho- logical traits were observed. Comparative Observations The deciduous upper canines of P; anubis. g; polyko— mgs and M; mulatta were similar in general outline. The canines of these three animals. however. could be distin- guished on the basis of the morphological details of the lingual surface (Fig. 3). The marginal ridge pattern was similar in all three animals; the mesial and distal ridges bordered the lingual surface and continued along the cervical 38 portion to some extent. The upper canine of P; anubis. how- ever. did not possess a well defined lingual ridge as did the other two animal groups. Instead. the area of the lin- gual surface. central to the marginal ridges. was smooth and convex. In addition. the cervical portions of the mar- ginal ridges blended into the convex central cervical por— tion of the lingual surface and did not produce a cervical ledge (Fig. 3. h). The upper canine of g; pglykomos more closely re- sembled that of M; mulatta than that of P; anubis. As in .M; mulatta. there was a lingual ridge present on the lingual surface of the canine of g; pplykomos (Fig. 3. i). This ridge. however. arose from the cervical ledge at a point approximately 1/4 of the tooth's length from the mesial sur- face and passed almost vertically to the apex of the tooth. The cervical ledge of the lingual surface was created by the continuation of the marginal ridges along the cervical portion of this surface (Fig. 3. i). This ledge. in g; _pplykomos. was undercut labially to a much greater extent than in either of the above two animals. In addition to the differences in the cervical ledge and the lingual ridge. the develOpment of a distal tubercle. present in most M; 39 mulatta and P; anubis. was not observed in any of the 16 g; gpolykomos specimens in which this surface was unworn. THE UPPER FIRST MOLAR Macaca mulatta The upper first deciduous molar 0f.fl; mulatta had five surfaces: mesial. distal. buccal. lingual. and occlu- sal. The mesial aspect presented two cuSps (paracone and protocone). the mesial marginal ridge. and an anterior' ledge directly mesial to the paracone. The cervical portion of the mesial surface Slanted disto-lingually (Fig. 3. j). The paracone and protocone diverged from the occlusal edge through the middle portion of the crown at which point they converged sharply to the cervix of the crown. The paracone was of greater height than the protocone. The anterior ledge was an elevation in the enamel at the termination of the paracone's mesial triangular ridge (Fig. 3. j). The mesial marginal ridge sloped cervically from the termina- tion of the triangular ridge of the protocone and mesio- buccally to the high point of the anterior ledge. At this 40 point. it joined with the triangular ridge of the paracone (Fig. 3. j). The distal aspect presented two cusps (metacone and hypocone) and the distal marginal ridge. The distal surface uniformly slanted mesially from the occlusal edge to the cervix of the crown and was rounded buccally and lingually. The buccal cusp was of greater height than the lingual cusp. The distal marginal ridge traveled disto-cervically from the hypocone triangular ridge and bucco-occlusally to the trian- gular ridge of the metacone; the distal marginal ridge was rounded and bowed cervically (Fig. 3. j). The buccal surface presented two well defined cuSps (paracone and metacone). a buccal developmental groove. a small anterior ledge. and a slight depression distal to the anterior ledge. The crown converged mesially and distally from the occlusal surface to the cervix of the tooth. The paracone was larger than the metacone mesio—distally but both were approximately equal in height. The buccal devel- opmental groove passed between the two cuSps. and ended in the middle third of the crown. The anterior ledge. as stated above. was an enamel enlargement at the mesial end of the paracone. A depression existed between this ledge and the ca: a: WGII at d1: the hy] 41 paracone; it extended from the occlusal surface to the middle portion of the crown. The lingual surface consisted of two cusps (protocone and hypocone) and a lingual developmental groove. The crown converged sharply mesially and distally from the occlusal surface to the cervix of the tooth. The two lingual cuSps were of approximately equal size in height and length. The mesial surface of the protocone leped mesio—buccally at a sharp angle until it approached the mesial marginal ridge; at this point it continued more gradually in a mesio-buccal direction. The lingual develOpmental groove continued onto the cervical third of the crown. The distal surface of the hypocone was smooth and gradually rounded. The occlusal surface was trapezoid in outline and had four cusps: paracone. protocone. metacone. and hypocone. The two mesial cuSps were united by a transverse ridge as were the two distal cuSps. and thus formed the bilOphodont molar pattern (Fig. 3. j). The relative position of the mesial cusps and the Shape and direction of the lOphodontS was variable in the Present sample. The arrangement of the cusps of the upper first molar could be absolutely determined in 76 of the 42 Macaque skulls. Of these skulls. 11 (five males. Six females) showed an arrangement in which the paracone and protocone were located directly opposite one another. In the remaining 65 Skulls (29 males. 36 females) the paracone was located further mesially than the protocone. The mesial lophodont. in this case. traveled from the protocone mesio-buccally to the paracone. The distal cuSps were always located Opposite one another. Three types of distal lophodonts. however. were observed. The first type consisted of a straight ridge which traveled from the tip of the hypocone directly buccally to the tip of the metacone (Fig. 3). This variety was observed in 72% of the males and 63% of the females. Type 2 consisted of a ridge which traveled buccally from the hypocone for a variably short distance. traveled distally. and continued buccally to the tip of the metacone. This variety was pres— ent in 4% of the males and 12% of the females. Type 3 con- sisted of a bifurcated. diamond shaped ridge with a depres- sion between its two parts. This variety existed in 24% of the males and 25% of the females. The occlusal surface presented a central fossa be— tween the mesial and distal lophodonts as well as a mesial 43 fossa and a distal fossa. The mesial and distal fossae were bounded by mesial and distal marginal ridges respectively (Fig. 3). A central developmental groove traveled from the central fossa mesially and distally between the two mesial and the two distal cusps. crossed the respective lOphodonts and ended in the mesial and distal fossae respectively. The course of this groove described a 'w' with the base of the letter located along the lingual surface (Fig 3. j). Buccal and lingual developmental grooves were also present on the occlusal surface. These grooves traveled be- tween the respective buccal and lingual cusps and continued onto the buccal and lingual surfaces respectively. The anterior continuation of the mesial triangular ridge of the paracone was noted above. It was continuous with the mesial marginal ridge and was reSponsible for the trapezoid shape of the occlusal surface. Comparative Observations Comparative observations were made difficult because of the high incidence of wear on the first maxillary molar of P; anubis. The general shapes of all five surfaces of this molar of P;_anubis were similar to the respective 44 surfaces of M; mulatta. lg; anubis also resembled M; mulatta in relative cuSp sizes. the marginal ridge forms. and de- velOpmental groove configurations. The relative cusp posi- tions and lOphodont forms and directions were variable as in M: mulatta. Unfortunately. the relative cusp arrange- ments could be determined in only four specimens. Two of these showed an arrangement of the mesial cusps. and the distal cusps. Opposite one another. In the remaining two casts. the buccal cuSps were located mesial to the respective lingual cuSps. The shape and direction of the lophodonts was observed in five specimens. In all but one of these. the lophodonts crossed from one lingual cusp to the opposite buccal cusp. In the one exception. the distal lophodont traveled mesio—lingually from the metacone and joined with the protocone. Enough specimens were not available to clearly define diagnostic features for P; anubis. In 9; pplykomos. the general outline of the five surfaces of the upper first molar were similar to the above two animals (Fig. 3. 1). However. several subtle yet con- sistent characteristics differentiated this molar of g; ‘polykomos from those of the above two Cercopithecinae species. First. the distal cusps were relatively smaller 45 in length and height than the mesial cusps. Secondly. the mesial triangular ridge of the paracone extended anteriorly but an anterior ledge was not present. Associated with this trait was the absence of depression on the buccal surface of the mesial portion of the paracone. (In one male specimen. a ledge and a depression were observed.) Thirdly. the mesial and distal marginal ridges traveled more cervically and the mesial and distal fossae were therefore less pronounced. The cusp arrangements in gppolykomos was observed in 12 specimens (five females; seven males). Three males and two females demonstrated an arrangement in which the lingual cuSps were located directly Opposite the buccal cuSps. In one male and one female. the paracone was located mesial to the protocone. In three males and two females. both buccal cuSps were located mesial to the respective lingual cuSps. The lophodont configuration was observed in 12 Specimens. Four males and three females demonstrated the lophodont M; mulatta type 1 arrangement. Two males and one female possessed a pattern similar to the type 2 arrange- ment Of.fl1 mulatta. One male and one female demonstrated a configuration in which the distal lophodont passed mesio- 1ingually from the metacone and connected with the protocone. 46 The above lophodont forms agree with Remane's description of variations in the upper first molar of Colobus (Remane 1951; 162). The latter two animals (and the one §;_anubis Specimen mentioned above) possessed Remane's 'Crista oblique' lOpho- dont pattern. It should be apparent that the upper first deciduous molars of the above three species were quite variable. Be- cause of this high degree of variability. it was difficult to clearly state descriptive diagnostic features and. there— fore. the non—metric features of this tooth did not provide good criteria for differentiating between animal groups. THE UPPER SECOND MOLAR Macaca mulatta The upper second deciduous molar. as Shown in the previous section. was larger than the first maxillary milk molar. The morphology of the second molar. however. did not differ greatly from that of the first deciduous maxil- lary molar. The crown had five surfaces: mesial. distal. buccal. lingual. and occlusal. As in the first deciduous 47 molar. the tooth had three roots. two buccal and one lingual. A groove was present on the lingual surface of the lingual root; it continued from the cervix of the crown to the apex of the root. The mesial aSpect presented two cusps (paracone and protocone). the mesial marginal ridge. and an anterior ledge. The paracone was slightly higher than the protocone. The mesial surface was trapezoid in outline due to the lingual slope of the protocone. The anterior ledge was rectangular in shape and leped distally at the cervical end. It re- ceived approximately equal extensions from the triangular ridges of the paracone and protocone. Because of this. the anterior ledge differed from that of the first upper molar which received no extension from the protocone. Related to this. the ledge ram: directly bucco—lingually; it did not slope lingually as in the first molar. The cervical line of this aspect sloped from buccal to lingual. The features of the distal surface were similar to those of the upper first molar. This surface presented two cusps (hypocone and metacone). and the distal marginal ridge. The distal surface was trapezoid in shape and leped mesio- Cranially. As in the upper first molar. the cervical line 48 sloped slightly lingually. the buccal cusp was higher than the lingual. the cervical portion of the hypocone leped mesio-lingually. and the distal marginal ridge was rounded and bowed cranially. The buccal surface presented two cusps (paracone and metacone). a buccal developmental groove. an anterior ledge. and a Slight depression distal to the anterior bulge. The buccal surface did not differ greatly from the buccal aSpect of the upper first molar. Both molars were similar in general shape. relative Sizes of the cusps. and the posi- tioning and extent of the grooves on the buccal surface. The only notable difference was the greater distance from the apex of the two buccal cuSps to the cervix in the second upper molar. The lingual surface of the upper second molar. how— ever. differed markedly from that of the upper first molar. Whereas the lingual suface of the first molar was almost triangular in outline. this surface of the second molar was trapezoid with the longest base at the occlusal edge. As in the upper first molar. both lingual cusps were of equal height. The hypocone of the second molar. however. was more rObust mesio-distally than that of the upper first molar; 49 a fact which was mainly responsible for the trapezoid shape of this surface. Neither cusp converged cervic- ally to as great a degree as in the upper first molar and the mesio-buccal slope of the protocone was not as sharp as that of the upper first molar. The two molars were similar in the positioning and extent of the lin- gual developmental groove. Unlike the first molar. however. a triangular pit was present mesial to the protocone; it was formed by a diagonal extension from the mesio—linguo-occlusal angle to the cervical third of the protocone. The occlusal surface was rectangular in Shape and presented four cuSps: paracone. protocone. metacone. and hypocone (Fig. 3. m). The two mesial cuSps were united by a transverse ridge as were the two distal cuSps (Fig. 3. m). The relative cusp positions were more stable than in the upper first molar. In all of the upper second molars observed. the mesial cuSps were positioned opposite one another as were the two distal cuSps. As in the first upper molar. the buccal cuSpS were higher than the lingual cuSps and the same apparent cusp size order prevailed. 50 The variations in the distal lophodont morphology were observed in the present sample of upper second molars; these three were the same as the three types defined above for the upper first molar. The frequency of occurrence for each type was calculated by studying 88 specimens which were in excellent condition. The percentages for types one. two. and three. were 55%. 27%. and 18% for the males and 55%. 30%. and 15% for the females respectively. As in the first molar. there were central. mesial. and distal fossae. These. again. were connected by a central developmental groove. Likewise. there were buccal and lin- gual develOpmental grooves which traveled between the re- spective buccal and lingual cusps (Fig. 3. m). Mesial and distal marginal ridges were also present. These traveled from the respective mesial and distal cuSps. peripherally to the mesial and distal fossae and formed the boundaries of the occlusal surface. The extension from the paracone supported a small elevation at its most mesio-buccal point. 51 Comparative Observations The upper second molars of Papio anubis and Colobus (polykomos resembled those of Macaca mulatta in the morphology of the mesial. distal. and buccal surfaces. In 9; polykomos. however. the second upper molar. as in the first molar. lacked an anterior ledge and the buccal surface therefore lacked a depression mesial to the paracone. The most dif- ferentiating characteristics. however. occurred in a consid- eration of the lingual and occlusal surfaces. In 3; anubis. the protocone was more robust than the hypocone (Fig. 3. n). This was the reverse of the condition noted for M; mulatta. In addition. the protocone extended further lingually than the hypocone; thus. the trigon's bucco— lingual dimension was greater than that of the talon. This observation was verified by taking the average of the dif- trigon — talon . ). For sample Size ferences in measurements of 37 teeth ( _P; anubis. the resulting average difference was .64 mm. For M. mulatta. however. the average difference was only .31 mm. (also based on 37 animals). To ascertain whether the above difference was due merely to dissimilarity in gross size. the ratio of the average difference to the average bucco-lingual average (trigon - talon) dlmen51on was calculated ( average greatest B-L measurement 52 X 100). For E; anubis. this ratio was 8.0; for M; mulatta. it was only 5.3. In addition to the above differences. the lingual surface of four Specimens of P; anubis (two males. two fe- males) demonstrated a tubercle between the two lingual cusps. This was not observed on any molar of M; mulatta nor 9; pg- lykomos. As in M; mulatta. the cuSp positioning and lophodont forms Of P; anubis and_gp_polykomos were more stable in the second upper molar than in the first. In the nine available casts of P; anubis. all second molars possessed a cuSp arrangement in which the paracone was located directly Oppo— site the protocone as were the two distal cuSpS. Likewise. the lophodonts of these nine specimens passed directly from one buccal cusp to the Opposite lingual cusp. This was also true for the 15 available Specimens Of S; polykomos (Six females; nine males). In addition to the differences of the buccal surface. _Q;prlykomos differed from the above two animals in degree Of develOpment of the lOphodontS and in the relative Sizes Of the lingual cuSpS. The lOphodontS in the g; polykomos upper second molar were very pronounced and were only Slightly 53 cut by the central develOpmental groove (Fig. 3. d). Secondly. the lingual cusps were of approximately equal height and mesio— distal length. The average trigon-talon difference was found to be .063 mm. The above ratio was found to be only 1.1. demonstrating the almost equal Size of the talon and trigon (Fig. 3). The Mandibular Dentition THE LOWER CENTRAL INCISOR Macaca mulatta The complex morphology and variability Of the upper central incisor were not noted for the mandibular first in— cisor. The lower central incisor was positioned vertically in its socket and had four surfaces: mesial. distal. labial. and lingual. The mesial and distal surfaces were triangular in Shape with the apex at the occlusal surface and were in- clined distally and mesially respectively. These two sur- faces were rounded in both directions. The labial surface was trapezoidal with the longest base at the incisal edge of 54 the tooth. The labial surface was smooth and rounded in both directions. The lingual surface was triangular with the apex at the cervix of the tooth. This surface Slanted lingually from the incisal edge to the cervix of the crown. The incisal third was flat and approximately rectangular in cross section; the cervical third was rounded Sharply and was approximately triangular in cross section (Fig. 4. a). In four partially erupted teeth of the Fort Johnson Specimens. Slight marginal ridges were noted but these disappeared before the tooth was completely erupted. In two newly erupted teeth. Slight in- dentations along the midline of the tooth were noted but these again soon disappeared. The root was cylindrical in the upper quarter of its length; in the apical third. it be- came flattened labio—lingually and hooked lingually at the apex. Comparative Observations The morpholOgical description of the lower central incisor 0f.fl; mulatta applied equally well for the lower first iJucisor of P; anubis (Fig. 4. b). The labial surface was more 55 rectangular in outline but. aside from metric differences. it was difficult to distinguish between the lower first in— cisors of the two groups of animals. The mandibular central incisor of g;_polykomos. how- ever. differed greatly from both of the above animals. The distal portion of the incisal edge Sloped disto-cervically giving both the labial and lingual surfaces a more rounded outline. The greatest difference appeared in the morphology of the lingual surface. The lingual surface was bordered by mesial and distal marginal ridges which were continuous with an incisal ridge and joined cervically to form a smooth curve. The lingual surface. central to the marginal ridges. was very convex and described a well defined lingual fossa (Fig. 4: C) . THE LOWER LATERAL INCISOR Macaca mulatta The mandibular lateral incisor was slanted in its Socket as was the upper second incisor. The crown of this tooth was also Similar in outline to that of the maxillary 56 second incisor. i.e. the incisal edge was slanted disto- cervically and the mesial surface was of greater height than the distal surface. The lower lateral incisor presented four surfaces: mesial. distal. labial. and lingual. The mesial and distal surfaces were triangular with the apex at the incisal edge. Both surfaces were Slanted distally and mesially respectively and they were rounded in both direc- tions. The cervical lines of both surfaces were inclined incisally from both the labial and lingual surfaces. The labial surface was triangular with the apex at the cervix Of the tooth. The surface was smooth and curved in both directions. There was a small depression located Slightly mesial to the distal surface which was positioned Opposite the triangular. lingual depression. The incisal edge described an elongated 'S.‘ i.e. there was a depression on the edge SO that approximately the distal third was lower than the mesial two—thirds (Fig. 4. d). The lingual surface. like the labial. was triangular in outline with the apex at the cervix of the tooth. A dis- tal marginal ridge. a Slight mesial marginal ridge and a Short incisal ridge were present on the lingual surface. All three ridges were poorly defined (Fig. 4. d). The 57 incisal ridge was present on only the distal portion of the incisal area. The mesial marginal ridge passed diagonally from the termination of the distal ridge to the mesio-incisal angle. The distal marginal ridge was continuous with the poorly defined mesial ridge and with the Short incisal ridge (Fig. 4. d). The distal and incisal ridges. and the disto- cervical portion of the mesial ridge. bordered a triangular lingual depression. The mesial portion of this lingual de- pression blended smoothly into the remainder of the lingual surface (Fig. 4. d). Comparative Observations Because of wear. Observations were possible on only four female Specimens of P; anubis. The lower lateral in- cisor of g; anubis was difficult to distinguish from that of M; mulatta. The labial surface was relatively longer meSiO-distally than in the lower Second incisor Of M; EE- latta and the ridges of the P; anubis lower lateral incisor were more poorly defined. Aside from these differences. the morphological features were Similar in both groups of animals (Fig. 4. e). 58 In 9; polykomos. the lateral incisor was caniniform. The incisal edge Sloped cervically in both directions pro- ducing essentially a one cuSped tooth. A tubercle. or ledge. was present at the termination of the distal SlOpe (Fig. 4. f). Mesial and distal marginal ridges were present on the lingual surface. These were continuous with a slight in- cisal ridge and a cervical ledge which SlOped meSiO-cervically. Thus. a rhomboidal. smooth and Slightly convex lingual depres- sion was outlined (Fig. 4. f). THE LOWER CANINE Macaca mulatta The lower canine had four surfaces: labial. lingual. mesial. and distal. Each of these surfaces was roughly tri- angular with the base at the cervix of the crown. The mesial surface was smooth and rounded in both directions. The cer— vical line of this surface Slanted incisally from labial to lingual. The surface inclined distally at the cervical third HOO Hmnuusw mmmm mmmwfln Hmcflmuma HmumHO USO Hmamma umcoo Imnmm co OmUOH Hoflnmucm 0s “mmmso HmflmmE SOS» HOHHmEm mmmso Hmumflo .ucmmmum OOOOH Hmsmsfla UOGHMOO Ham? aucmmmum OOOHH HOSOSHH HOOHOHO> “maouonsu HmumHO OS “HOOHHOOEESO HOHSOGOHHB .ucmmmum OOUOH HmSSCSH smcnmma Ham; “woven Hmsmsfla Hmsommflw OS “OommuSm Hmmflosfl um xwmm mmmnm Haasmcmflne .momm IMSm Hmsmcfla O>mocoo “OmUOH HOOH>HOO mmnmq .muumase 4m.ou HOHHEHm .OOOMHSO Hmsm ISHH xm>soo Ucm SDOOEm uucmmnm mmOHH Hmso ImmHO Hmsmsfla uucmmwum maoumnsu HmumHO “HMO IHHDOEE>OGD HMHSOSOHHB .OmUHH Hmco IOOHO uuonm “xfl>nmo um xmmm Ommnm amasmsmflue .haam ImHOSH mocmumflw uuonm maso mcflvcmuxm OmOHH Hmsmsfla .mmmnm OHOQEOQm .cumuumm OOOOOSQOH HmumHO USO COHUHOOQ mmso HmHmOE OHQMHHO> “mcoomumm co usmmmum OmOmH Hmflmmfi “Osouomhn .choomuwe .Osooououmv .Osoomumm "ONHm mmsu .maamflmmE OSHBOQ USO ucwmmum OmOHH Hmco ImmHU Hmsmsfla “usmmmum OHOHOQSO HmumHO “HMO Ifluumeemmcs HMHDOQOHHB .OmOHH Hmsmcfla Hmso ImmHO mcoa uxH>HOO um xmmm Ommnm “masmsmflua .OmOO HmmHOSH Hmmc Ocflvcmuxm OmOHH Hmsmcfla Hmsommflp mo mocmsqmum 30H: “Ommnm HOHSOCOHHB SOHO: umuflm xxuwmmb OSHSMU «Hmmmb uOmHOSH Hmumumq summmb HomHOSH Hmupcmo RHOQQD moonwaom,mSQoHOU mflnscw oflmmm MUHMHDE MONUME Snooe HOEHS< 76 .mmmum Omoam HmHmOE Lucmmmum OOOHH Hmsmsfla Hmsommflc “Dam Immnm mmmOHH amcfimumz .OmOOH HMOH>HOO HOSOOHH USO mOmUHH OOQOHO>OO HHOB “OmOO Hmmflosfl um xmmm Smasmsmfluu .Euowflcficmu .mmmom Hmsmsfla UOOAMOU HHOB uucmmmum mmOHH Hmmflocfl “USO Immnm mmmOHH Hmsflmumz .mmwso Hmomcfla cmm3umn usmmmum HO>OS OHOHOQSD “Amcooououm on Hmsqm usoflm Ocooommmv conu ou HOSUO OOOEHO com Ifluu “Ocoomumm co USO Immum mmOOH HmfimOE Oz .mupmHDE quou HOHHEHm .OOQOHO>OO hauoom DOD ucmmmum mOmOHH HmchHmE umeO Hmmflo IcH Ommmnm.m. uxfl>umo um xmmm HOHSOSMHHB .muumHSE 4w.ou HOHHEHm .mmmso HmsmcHH SOOBDOQ ucmmmum mOEHuOEOO OHO lumflsu “Amcooououm awry HOHHmEm £058 Ocooomxmv SOHmu SOLO ummuma LOSE comfluu umcoomumm so ucmmmum OOOOH HOHOOE .Hmsvmum Omoam HmHmOE “OOOHH amsmcfla o: “UGO Immum momUHH Hmcfimumz .OOQOHO>OO hamumHOOOE Osm ucmmmum mmmwflu Hmcflmuma "OOOO Hmmflo ISH Ommm£m_m. “xH>HOO um xmmm Smasmcmflue .uuom mam mo mmmOHH OS OOOMHSO SpooEm uxfl>umu um xmmm HOHSOSOHHB .mmmso Hmsm ISHH SOOBOOQ ucmmmum um>mc OHOHOQSO “Amcoo lououm SOLO HOHHmEm mcooomxmv conu Smnu q-m ”magma Saunmaam somfluu “msoomnmm so ucmmmum OmOOH Hmflmmz OSHSOU «HOBOA HomflocH Hmnmumq «HOBOA HomflOSH Hmuucmo SHO3OA SOHO: Ocoomm Seaman: onoxNAQm.msnoHoo manscm oflmmm. muumase momomz Snooe Hmeflcm 77 .mommusm HmmSHOOo¥« .maso OOMMHSO HOSOOHQS .mmOflH HOOHOHOE HmHmOE Ommmnm OHHOQOHOQ “Acoflu loanumsoo HOHOOEV mmmso HmumHO can» Hmnumm Iou ummoHo Omcofluflmom mmmsu HMHOOE “mucowonm IOH Ocm mmmso OOSHMOO HHO3 “mmmso HOOOSQ 030 Cmm3umfl usmmmum Hm>mc OHOHOQSO uucmwmmm HO>OO ESHOOEHODSH ESHSOHOQSB .mSSmHSS 4m.ou Hmaneflm .OmOHu Hmcflmnms HOHOOE Ommmflm D “mmmso HmumHO mm Uwsofluflmom mmmoo HOHOOE “mucowonm IOQQOH Ocm mmmso OOGHM IOU mamumnmwoe “mmmso Hmoosfl 03» SOO3uOQ uqm Immum mOeHuOEOO OHOHOQ In» uucmmmnm haucmsvmum ESHOOEHOOCH ESHSOHOQSB .mmmmwmm..z 0» Hmfiflsflm .OOOHH HmchHmE HOHOOE Ommmnm D “mmmso Hmumflv Op HOHHEHO Umcoflufimom mmmso Hmfimme amuSOO Ionmoa USO mmmso Omcflm IOO >HmumuOOOE “mmmso HOOOSQ OB» Smm3umn usmmmum mmEHuOEom OHO lumnsu uucmmmum hamumn Esflcmfinmusa ESHDOHOQSB .OODOHOOH mmmom HmHmOE Lucmmmum maonoflsu Hoflumucm “umnuocm Ono mnemommo mmmso Hmumflw “OHSOO Iououm ou HmumHO OHSOO ImuOE “Ommmnm ummmflam HMHOE UCOUmW ¥¥Hm301H SOHO: amuse «SHOBOA moonmaom,mSQoHOU manscm oflmmm MUHMHSE MUMUME SDOOB Hmeflca 78 separated from g; polykomos. It is suggested. therefore. that morphology Should be the basis for the generic designa- tion of a Specimen. Before this can be unequivocally stated. however. considerable work remains to be conducted on related Species of the above animals. Given the genus of a Specimen. the species may be diagnosed if sufficient data on metrics are available from previous studies. Hence. the Species of a Specimen of Papio and Macaca might be determined from the comparative metric study Of Biggerstaff and the comparison of two Species Of baboon in the present work. Further evidence of the value of metrics in determining species level is indicated in the study of two groups Of Cercopithecus aethiops sabaeus by Ashton and Zuckerman (1950b. 227). The two groups of Cerco- pithecus were separated for as many as 75 to 100 generations. Ashton and Zuckerman Observed that the morphology of the two groups had remained constant while the St. Kitts monkeys had an increase in tooth dimensions (Ashton and Zuckerman 1950b; 227). This suggests that in the formation of new Species the ciimensions of teeth are more subject to change while morpho— logical characteristics remain relatively constant. It is Obvious that much additional information is needed to 79 determine the accuracy with which Species may be designated on the basis of deciduous tooth measurements. Two further topics deserve mention before concluding. First. the functional and adaptive Significance Of metric and morpholOgical deciduous dental traits will be considered. The adaptiveness of the former trait is relatively easily understood. If. in the course of speciation. larger animals with larger permanent dentitions are positively selected for. increased Size of the deciduous dentition will result and will function to fulfill the increased nutritional needs of the larger juveniles. The author feels that if selective forces have acted on the dentition itself. these forces were directed toward the permanent teeth rather than toward the deciduous (although forces acting directly on the decid- uous dentition are not inconceivable). The morpholOgical traits are more varied and are therefore more difficult to understand. The deciduous teeth must serve the juvenile macaque for approximately three years (Hurme and van Wagenen 1953. 310; Schultz 1933; 23) and the length of serviceable time for the milk teeth of other Old World monkeys Should be comparable. Throughout this period. 80 the dentition Of the juvenile must function in the same eco- logical System as that of the adult. AS Gregory pointed out. the food of the young is Often the same as the adult and an efficient crown pattern is therefore important in both sets Of teeth (Grengory 1922; 470). In general. the morphology of the deciduous teeth should resemble that of the adult. AS noted above. this is especially true for the second de- ciduous molars which have closely paralleled the first perm- anent molars in their evolutionary development. It appears that. in the course of evolution. as the adult dentition be- comes adapted to a Specific ecological niche. the deciduous dentition follows with Similar traits which will insure the survival of the juveniles. The functional and adaptive sig— nificance of a morphological trait Of a deciduous tooth. as with the permanent teeth. must be interpreted in view of the particular habitat of the Species. Discussion of the adaptive and functional Signifi- cance of Specific traits is difficult and. at the present time. a number of unanswered questions can merely be raised. What are the functions of the incisors' lingual fossa. the caniniform mandibular lateral incisor or the mesial constric- tion of the mandibular second molar of the leaf eating 9; 81 polykomos? Likewise. is there any functional advantage in comparable traits in the omnivorous M; mulatta and P;_anubis? Or are such traits merely artifacts of evolution? Several Of these questions may be answered when the extent of these morphological traits have been determined in animals Of Similar habitat. Secondly. it has Often been stated that the deciduous dentition retains primitive features and that examination of these teeth will reveal primitive characteristics of ances- tral forms. This belief was the principle guide of Remane's search for evidence for the origin of bilOphodontism in the Cercopithecidae (Remane 1951; 162). JOrgensen. starting with a firm comparative foundation. listed 33 traits which he con- sidered to be primitive (conservative) in Danish deciduous teeth (J¢rgensen 1956. 180—183). Gregory pointed out that the deciduous dentition need not be strictly primitive but may also possess adaptive (progressive) traits (Gregory 1922. 470). More recently. this view has been expressed by von Koenigswald (1967. 779). The question now raised is which traits of the de- Ciduous teeth should be considered progressive and which 82 should be classed conservative. In general. a caniniform lateral incisor would be considered primitive (Jorgensen 1956. 177). Should the lateral incisors of ngpolykomos be considred primitive? Caniniform lateral incisors happen to appear in the adult dentition as well as in the milk teeth Of E; polykomos. The caniniform deciduous lateral incisor may be a development parallel to that of the permanent lateral incisors and a secondary adaptation in both sets. Problems also arise in considering the 'crista obliqua' observed by Remane and in the first upper molar of g; polykomos. The majority of first maxillary molars do not demonstrate this trait. The author feels that such anomolous traits are noteworthy but should not be considered primitive nor Should evolutionary changes be inferred from them. On the basis Of present knowledge of the extent of morphological traits in the deciduous dentition of the Old ‘WOrld monkeys. it is difficult to determine which traits Should be considered primitive. The author agrees in part \vith von Koengswald when he stated that as the deciduous dentition of the Cercopithecidae adapt to new conditions they are becoming 'progressive' and retain no traces of earlier stages (von Koengswald 1967; 782). If broad 83 comparative data on the deciduous dentitions of all living primates were available. however. the degree of divergence of genera from the 'ancestral type' may be determined. In this sense. the 'primitiveness' of certain generic traits may be defined. The final resolution of the primitiveness question must. of course. lie in the discovery of actual ancestral fossil forms. S UMMARY Odontometric and comparative morphological studies were conducted on the deciduous dentition of the following Old World monkeys: 150 Macaca mulatta; 20 Papio anubis; l7 Colobusmpolykomos. Comparative statistical analysis of tooth measure- ments from these animals revealed: the presence of metric bilateral symmetry in all teeth of the above three genera; the appearance of sexual dimorphism in the deciduous molars of all three genera; intraspecific variations Of each Species; interspecific differences for Macaca and Papio; statistically significant differences between animals of the three genera considered. Detailed morphological and comparative observations \nere presented for all three animal groups. ML mulatta closely resembled E; anubis in many morphological features. \Mhereas both groups differed from the morphology of g; jgglykomos. Distinguishing features for each group were Summarized and diagnostic criteria were suggested. 84 APPEND IX 86 TABLE l.—-Correlations of right and left Side dentitions within each Species. Macaca mulatta Mesio-distal measurement Tooth N Mean r Tooth N Mean r rii 121 4.817 .847 ril 136 2.985 .831 1i 125 4.754 1il 134 2.995 .2 r12 124 3.861 .745 ri2 137 3.056 .708 1i 114 3.778 1i2 131 3.152 rci 139 4.588 .809 rcl 140 4.114 .806 1c 132 4.597 lcl 142 4.160 rmi 151 5.907 .901 rml 147 6.295 .908 1m 151 5.905 1ml 148 6.291 rm: 151 6.498 .939 rm2 150 6.594 .961 lm 151 6.490 1m2 150 6.551 Bucco—lingual measurement rii 120 3.683 .799 ril 135 2.975 .893 1i 126 3.705 lil 134 2.963 rig 124 3.063 .871 ri2 137 2.804 .768 1i 114 3.066 1i2 131 2.727 rci 139 3.664 .884 rcl 140 2.916 .772 1c 132 3.661 1cl 142 2.896 rm: 151 5.138 .939 rml 150 3.969 .886 lm 151 5.114 1ml 150 3.978 rm: 151 5.827 .954 rm2 150 4.847 .932 lm 151 5.837 1m2 150 4.837 87 TABLE 1 (continued) Papio anubis MesiO-distal measurement Tooth N Mean r Tooth N Mean r ri: 11 7.073 .955 :11 11 4.736 .725 1i 11 6.982 111 12 4.783 2 ri2 12 6.050 .943 ri2 11 5.109 .877 11 9 6.056 112 12 4.983 rci 17 6.635 .923 rCl 18 5.867 .936 1C 19 6.568 1Cl 16 5.981 rm: 18 8.028 .907 rml 20 8.745 .894 1m 16 7.919 lml 19 8.795 rm: 19 8.805 .888 rm2 19 8.937 .942 1m 18 8.789 lm2 19 8.868 Bucco—lingual measurement ri: 11 4.827 .952 r11 11 4.109 .950 11 11 4.882 111 12 4.000 ri: 12 4.542 .959 r12 11 4.227 .877 11 9 4.644 112 12 4.108 rc: 17 4.671 .913 rcl 18 4.094 .874 1c 19 4.747 1cl 16 4.012 rm: 18 6.489 .906 rml 20 5.220 .958 1m 16 6.569 1ml 19 5.310 2 rm2 18 8.039 .956 rm 19 6.847 .960 1m 19 8.084 1m 19 6.921 88 TABLE 1 (continued) Colobus polykomos MeSiO—distal measurement Tooth N Mean r Tooth N Mean r rii 11 3.809 .884 r11 12 2.458 .702 1i 11 3.727 1il 10 2.470 ri: 13 3.369 .798 ri2 12 3.033 .842 1i 11 3.427 1i2 11 3.100 rci 15 4.973 .909 rcl 15 4.453 .882 1c 15 5.013 1cl 14 4.407 rm: 17 5.271 .942 rml 16 5.744 .821 1m 17 5.247 1ml 17 5.765 rm: 16 6.419 .866 rm2 16 6.537 .917 1m 16 6.412 1m2 15 6.520 Bucco-lingual measurement ri: 11 2.754 .973 r11 12 2.300 .843 11 11 2.764 111 10 2.310 rig 13 2.592 .859 ri2 12 2.525 .860 11 11 2.609 112 11 2.554 rci 15 3.247 .927 rcl 15 2.913 .768 1C 15 3.287 1C1 14 2.936 rm: 17 4.323 .763 rml 16 3.387 .739 1m 17 4.329 1ml 17 3.382 rm: 16 5.500 .805 rm2 16 4.781 .885 1m 15 5.520 1m 16 4.787 2 89 TABLE 2.—-Comparison of male and female tooth measurements within each species. Macaca mulatta Measure— Sex Tooth N Mean S.D. C.V. S.E. t ment M ri: M-D 57 4.817 .331 6.87% .044 .007 F ri M-D 64 4.817 .245 5.09 .031 M r1: M-D 63 3.824 .227 5.94 .028 -l.858 F ri M—D 61 3.900 .229 5.87 .029 M rci M-D 71 4.572 .235 5.14 .028 -.890 F rc M-D 68 4.604 .193 4.19 .023 M rm: M-D 75 5.968 .259 4.34 .030 2.731 F rm M-D 76 5.846 .288 4.93 .033 M rm; M-D 64 6.593 .289 4.38 .033 3.878 F rm M-D 72 6.404 .311 4.86 .036 M ril M-D 64 3.011 .216 7.17 .027 1.449 F ril M-D 72 2.962 .173 5.84 .020 M ri2 M-D 68 3.038 .253 8.33 .031 -.758 F ri2 M-D 69 3.074 .296 9.63 .036 M rcl M-D 70 4.134 .232 5.61 .028 1.056 F rcl M-D 70 4.094 .216 5.28 .026 M rml M-D 73 6.379 .274 4.29 .032 3.454 F rml M—D 74 6.212 .312 5.02 .036 M rm2 M-D 75 6.688 .307 4.59 .035 3.375 F rm M-D 75 6.500 .372 5.72 .043 90 TABLE 2 (continued) Macaca mulatta Sex Tooth Measure" N Mean S.D. c.v. S.E. t ment M r1: B-L 56 3.700 .155 4.19% .021 1.031 F ri B-L 64 3.669 .174 4.74 .022 M r1: B-L 63 3.068 .208 6.78 .026 .313 F ri B-L 61 3.057 .177 5.79 .023 M rci B—L 71 3.068 .205 6.68 .024 -.468 F rc B-L 68 3 057 .190 6.21 .023 M rm: B-L 75 5.180 .235 4.54 .027 2.313 F rm B-L 76 5.096 .211 4.14 .024 M rm: B-L 75 5.901 .234 3.96 .027 3.726 F rm B-L 76 5.754 .252 4.38 .029 M r1l B-L 63 2.967 .167 5.63 .021 —.541 F ril B—L 72 2.982 .160 5.37 .019 M ri2 B-L 68 2.798 .190 6.79 .023 -.305 F r12 B-L 69 2.809 .199 7.08 .024 M rcl B-L 70 2.901 .200 6.89 .024 —.902 F rcl B—L 70 2.931 .193 6.58 .023 M rml B-L 75 4.015 .206 5.13 .024 2.676 F rml B-L 75 3.924 .208 5.30 .024 M rm2 B-L 75 4.900 .221 4.51 .026 2.971 F rm B-L 75 4.795 .213 4.44 .025 TABLE 2 Sex Tooth (continued) Measure- 91 Papio anubis S.D° .331 .545 .227 .617 .365 .417 .415 .429 .227 .453 7.77% .331 .172 .227 .186 .163 .120 .169 .124 .080 .136 1.208 -.423 .691 1.922 2.910 M ri: F ri .2 M r12 F ri M rci F rc M rm F rm 2 M rm2 F rm M . F :11 1 i: :2 2 M F :21 l M r F r21 1 M rm2 F rm N Mean ment M-D 1 7.700 M-D 10 7.010 M—D 1 5.800 M—D 11 6.073 M—D 5 6.740 M—D 12 6.592 M-D 6 8.300 M-D 12 7.892 M—D 8 9.100 M-D 11 8.591 M—D 1 5.000 M-D 10 4.710 M-D 1 5.200 M-D 10 5.100 M-D 6 5.900 M—D 12 5.850 M-D 8 8.800 M-D 12 8.708 M-D 8 9.012 M—D 11 8.882 .216 .273 .253 .469 .322 .363 .421 .500 .587 .595 .216 .086 .253 .148 .131 .105 .149 .057 .207 .179 .203 .285 .427 .476 92 TABLE 2 (continued) Papio anubis Sex Tooth Measure" N Mean S.D. c.v. S.E. t ment M ri: B-L 1 4.100 .155 -- .155 -3.005 F ri B—L 10 4.900 .254 5.18% .080 M r1: B—L 1 4.100 .208 —— .208 -1.101 F ri B—L 11 4.582 .419 9.14 .126 M rc: B—L 5 4.780 .217 4.54 .097 1.070 F rc B—L 12 4.625 .290 6.27 .084 M rm: B-L 6 6.767 .320 4.73 .131 2.012 F rm B—L 12 6.350 .450 7.09 .130 M rm: B-L 7 8.471 .269 3.17 .102 3.551 F rm B-L 11 7.764 .478 6.16 .144 M ril B—L 1 3.900 .167 -- .167 -.775 F ril B-L 10 4.130 .283 6.85 .089 M ri2 B-L 1 4.300 .190 -— .190 .247 F ri2 B-L 10 4.220 .308 7.30 .097 M rel B-L 6 4.267 .372 8.72 .152 “1.539 F rcl B-L 12 4.008 .317 7.91 .091 M rml B-L 8 5.425 .471 8.68 .166 1.783 F rml B-L 12 5.083 .383 7.53 .111 M rm2 B-L 8 7 212 436 6.04 154 2 931 F rm B-L 11 6 582 481 7 31 145 93 .045 .853 .495 .205 .293 .253 .598 .753 TABLE 2 (continued) Colobus pplykomos Sex Tooth Measure" N Mean S.D. c.v. S.E. ment M ri M—D 8 3.812 .368 9.65% .130 F ri M-D 3 3.800 .529 13.92 .305 M ri: M—D 9 3.333 .240 7.20 .080 - F ri M—D 4 3.450 .191 5.54 .095 M rc M-D 10 4.850 .310 6.39 .098 —2 F rc M-D 5 5.220 .148 2.84 .066 M rm: M-D 10 5.260 .217 4.12 .069 - F rm M—D 7 5.286 .302 5.71 .114 M rm: M—D 10 6 350 135 2.12 .043 —1. F rm M—D 6 6.533 .273 4.18 .111 M ril M-D 8 2 462 151 6.13% .053 F ril M—D 4 2 450 129 5.26 .064 M ri2 M—D 8 3.012 .247 8.20 .087 — F ri2 M—D 4 3.075 .512 16.65 .256 M rcl M-D 9 4.322 .268 6.20 .089 -2 F rcl M—D 6 4.650 .288 6.19 .118 M rml M-D 9 5.711 .242 4.24 .081 - F rml M-D 7 5.786 .254 4.39 .096 M rm2 M-D 10 6.430 .189 2.94 .060 -2 F rm M—D 6 6.717 .223 3.32 .091 94 TABLE 2 (continued) m Colobus_po1ykomos Sex Tooth measure- N Mean S.D. c.v. S.E. t ment M r1: B—L 8 2.737 .261 9.53% .092 —.377 F ri B-L 3 2.800 .173 6.18 .100 M Ii: B-L 9 2.567 .100 3.89 .033 -1.115 F ri B—L 4 2.650 .173 6.53 .086 M rci B-L 10 3.190 .256 8.02 .081 -l.235 F rc B-L 5 3.360 .241 7.17 .108 M rm: B-l 10 4.290 .238 5.55 .075 -.774 F rm B-L 7 4.371 .170 3.89 .064 M rm; B-L 10 5.470 .275 5.03 .087 -.612 F rm B-L 6 5.550 .207 3.73 .084 M r11 B-L 8 2.275 .183 8.04 .065 -.641 F ril B-L 4 2.350 .208 8.85 .104 M ri2 B-L 8 2.487 .173 6.96 .061 —.848 F ri2 B—L 4 2.600 .294 11.31 .147 M rcl B-L 9 2.900 .245 8.45 .082 —.274 F rcl B-L 6 2.933 .207 7.06 .084 M rml B-L 9 3.378 .083 2.46 .028 -.394 F rml B-L 7 3.400 .141 4.15 .053 M rm2 B-L 10 4.640 .107 2.31 .034 4.357 F rm B—L 6 5.017 .240 4.78 .098 95 TABLE 3.--Comparison of tooth measurements of Macaca mulatta. Papio anubis. and Colobus polykomos. Mesio-distal measurement Source of Sex Tooth . . S.S. d M.S F* ___ Variation M ril Among groups 15.99 2 7.99 71.01 Within groups 7.09 63 .11 F ril Among groups 46.56 2 23.28 246.10 Within groups 7.00 74 .09 M ri2 Among groups 5.99 2 2.99 57.36 2 Within groups 3.65 70 .05 F ri Among groups 46.31 2 23.16 239.03 Within groups 7.07 73 .10 M rcl Among groups 22.12 2 11.06 174.51 Within groups 5.26 83 .06 . F rcl Among groups 40.76 2 20.38 370.85 Within groups 4.51 82 .05 M rml Among groups 37.12 2 18.56 260.57 1 Within groups 6.28 88 .07 ' F rm Among groups 47.93 2 23.97 250.38 Within groups 8.81 92 .10 M rm2 Among groups 47.52 2 23.76 319.57 2 Within groups 6.69 90 .07 F rm Among groups 46.08 2 23.04 214.41 Within groups 9.67 90 .11 M ril Among groups 6.28 2 3.14 70.92 Within groups 3.10 70 .04 F ril Among groups 28.82 2 14.41 420.01 Within groups 2.85 83 .03 M ri2 Among groups 4.63 2 2.31 36.37 Within groups 4.71 74 .06 F ri Among groups 36.10 2 18.05 165.60 2 . . Wlthln groups 8.72 80 .ll 96 TABLE 3 (continued) Mesio-distal measurement Sex Tooth sonice.°f 5.8. d M.S. F* Variation M rcl Among groups 17.25 2 8.62 146.92 Within groups 4.81 82 .06 F rcl Among groups 32.08 2 16.04 268.21 Within groups 5.08 85 .06 M rml Among groups 48.91 2 24.46 299.34 Within groups 7.11 87 .08 F rml Among groups 68.22 2 34.11 299.90 Within groups 10.24 90 .11 M rm2 Among groups 41.13 2 20.57 190.26 Within groups 9.73 90 .11 F rm2 Among groups 54.46 2 27.23 173.06 Within groups 14.01 89 .16 Bucco—lingual measurement M ril Among groups 6.75 2 3.38 116.36 1 Within groups 1.80 62 .03 F ri Among groups 16.20 2 8.10 234.38 Within groups 2.56 74 .04 M ri2 Among groups 3.16 2 1.58 40.16 2 Within groups 2.76 70 .04 F ri Among groups 23.21 2 11.61 226.80 Within groups 3.74 73 .05 M rcl Among groups 8.48 2 4.24 94.78 1 Within groups 3.71 83 .05 F rc Among groups 10.24 2 5.12 117.52 Within groups 3.57 82 .04 A.)l._.- 97 TABLE 3 (continued) Bucco-lingual measurement Sex Tooth soche.°f 8.5. d. M.S F* __ Variation M rm1 Among groups 23.02 2 11.51 198.53 1 Within groups 5.10 88 .06 F rm Among groups 21.50 2 10.75 172.48 Within groups 5.73 92 .06 2 M rm Among groups 46.06 2 23.03 396.87 2 Within groups 5.16 89 .06 F rm Among groups 40.00 2 20.00 248.23 Within groups 7.25 90 .08 M ril Among groups 4.40 2 2.20 76.94 Within groups 1.97 69 .03 F ril Among groups 13.85 2 6.92 214.60 Within groups 2.68 83 .03 M ri2 Among groups 3.02 2 1.51 42.29 Within groups 2.64 74 .04 F ri2 Among groups 17.97 2 8.98 188.60 Within groups 3.81 80 .05 M rcl Among groups 10.40 2 5.20 108.10 Within groups 3.94 82 .05 F rcl Among groups 12.02 2 6.01 131.16 Within groups 3.89 85 .05 M rml Among groups 19.23 2 9.61 179.59 Within groups 4.76 89 .05 F rml Among groups 16.93 2 8.47 155.55 Within groups 4.95 91 .05 M rm2 Among groups 40.74 2 20.37 362.82 Within groups 5.05 90 .06 F rm2 Among groups 30.64 2 15.32 228.67 Within groups 5.96 89 .07 K . *Significance probabilities for all F statistics were (.0005. 953 TABLE 4.-—Relat1ve 51205 of teeth w1tn1n each Spcc1es. REEATIVE MESIO-DISTAL DIMENSIONS — 7 malo- m7 V m1 3 1l * c1 P 12 male— m2 > ml > c1 > 12 > il _ (4.3) (1.15) (.23) (.75) (.31) (2.3) (1.1) (.03) Macaca mulatta _ 2 . l . 1 _ 1 . 2 - . . temale- m 1 m I 1 ’ c ‘ 1 iemalc- 1112 > 1111 > C1 > 12 > 11 (.6) (1.0) (.2) (.7) (.3) (2.1) (1.0) (.1) * *_ male- m2 > m1 > 11 P c1 \ 12 male— m2 > ml > c1 > 12* > i1* (.8) (.6) (1.0) (.9) (.2) (2.9) (.7) (.4) Papio anubis 2 l l l 2 femalc- m > m > i > c 5 1 female— m2 > ml > C1 > 12 > i1 ( 7) ( 9) ( 4) ( 5) (.2) (2.9) (.8) (.4) 1 .1 .2 l . — / 3 > .. - > > > > ' male m(l 4)m(1 3)1(.i)1(.4)C mile 1112 1111 C1 12 11 p (.5) (3.3) (1.0) (.4) apio ur51nus 2 l 1 2 l (Freedman '62) female— m > m > i i i i c female- m > m > c > i > i 1.2 1.4 . . 2 l 1 2 1 ( ’ ‘ ) ( ’ ( ’ (.5) (3.0) (.7) (.9) male- m2 > m1 > c1 > 11 > 12 male- m > 1111 > C1 > 12 > 11 (1.1) (.4) (1.0) (.48) (.7) (1.4) (1.3) (.6) Colobus EEAXEEEQE female- 1112 > m1 > c1 > i1 > 12 female— m2 > ml > c1 > i2 > i1 (1.3) (.07) (1.4) (.4) (.9) (1.1) (1.6) (.6) RELATIVE BUCCO—LINGUAL DIMENSIONS - male- m2 > m1 > 11 > c1 = 12 male- m2 > 1111 > 11 > c1 > 12 (.6) (1.5) (.6) (.9) (1.1) (.07) (.1) Macaca mulatta 2 l 1 1 2 female- m > m > i > c = 1 female- 1112 > 1111 > il > C1 > i2 (.7) (1.4) (.6) (.9) (.9) (.05) (.1) male- m2 > m1 > c1 > 11* = 12* male- m2 > 1111 > i; > ci > i1. - . (1.7) (2.0) (.7) (1.8) (1.1) (.03) (.4) Papio anubis 2 l l 1 2 female- m > m > i > c > 1 female- m > ml > 12 > 11 > c1 (1.4) (1.5) (.3) (.04) (1.5) (.9) (.09) (.12) male- 11)2 > m1 > 11 > c1 > 12 male- 1112 > 1111 > C1 = 12 > il Pa 10 ursinus (1.3) (2.0) (.1) (.1) (2.0) (.5) (.1) (Freedman '62) female— mil>5)mtl ;)i% 3):? i)12 female- (112 > m > i > c1 = i2 ' ' ' (1.9) (.6) (.2) male- m > m1 > c1 > i1 > i2 male- 1112 > 1111 > C1 > 12 > 11 Colobus (1-2) (1-1) 1-5) (.2) (1.3) (.5) (.4) (.2) 221152E2§ female- m2 > m1 > c1 > i1 > 12 female- m2 > 1111 > c1 > 12 > 11 (1.2) (1.0) (.6) (.2) (1.6) (.5) (.3) (.3) *Based on small sample. 99 ._o>o_ oucon_ucou mo.ofl ozu um mac—«L: o_mum vcu «_naco m_mam c. mac—mcoe_v suOOu _oum_v|o_nue mo com_LooEOU ._ .m_u N! _E _u N. __ NE _E _u N. _— 0 use I I-.) . —O_0.W_ I m _ A moxvm voc_oEou I < .4 _ ANmm_ coEvumLu T m axe—mu: 0_ on u m . A>vaum ucomotav _ _m_..|€clo 9mm... 1 < .4 o . _ . + . _ . _ ..o . m < + _ - Y a < _ . . _ _ w h _ (N _ + _ 1 _ — é .if _ I. _ . . . . _ + . . _ (m . _ n 1 _ _ i _ _ _ _ . . + . _ _ lo- . . < _ o < _ n sun-w) 1 l (u 100 o<_-II. «oxen voc_aeou I :8. 368: > 32%).. 2 .. a . 3%.»?J . < :26. 85228 3.8.. 9: on aqua.“ v5 543:3 c. .8355... 58” 32.7803 3 83.393 .N .m: «E _E —u N. __ «E _E _o N. —~ _ . . _ + + 4 n . . i. + + _ _ . _ a < a < _ a < _ +* + l. _+ + + . _ . . _ _ _ _ 1 a _ u o < _ a < _ ._ _ n _ 1 0-“50 IIIID _ _ _ To Haw) l l (u 101 LINGUAL VIEW OF THE ANTERIOR TEETH m—d Macaca mulatta ’ Papio anubis Colobus polykomos Upper A c Central Incisor Upper D F Lateral lncisor Upper G H Canine OCCLUSAL VIEW OF THE POSTERIOR TEETH Macaca mule/Ia Papio anubis Upper First Molar Upper Second Molar fig. 3 MORPHOLOGY OF THE MAXILLARY DECIDUOUS DENTITION 102 LINGUAL VIEW OF THE ANTERIOR TEETH rn Macaca mu/affa Pap/o anubis Colobus polykomos Lower Central lncisor Lower Laleral Incisor Lower Canine OCCLUSAL VIEW OF THE POSTERIOR TEETH )—|-—b Macaca mulatta Pap/a any bi: Colobus polykomos Lower Fhsl Moku' Lower Second Nkflar “94 MORPHOLOGY OF THE MANDIBULAR DECIDUOUS DENTITION L ITERATURE C ITED Ashton. 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