93 Palaeont. afr., 14. 93-111. 1972 THE PARAPAPIO SPECIES OF STERKFONTEIN, TRANSVAAL, SOUTH AFRICA by L. Freedman & N. S. Stenhouse (Department of Anatomy and Raine Medical Statistics Unit, University of Western Australia, Nedlands, W.A. 6009) ABSTRACT The premolar and molar dimensions of the Parapapio specimens hitherto described from the South African lower Pleistocene deposits at Sterkfontein have been re-examined. The relatively minor sexual dimorphism in these teeth (except P3 length) was confirmed. 103 new cercopithecoid specimens from Sterkfontein were tentatively assigned to the 3 previously described Parapapio spp. from that site. The new and old samples of each species were compared and then pooled. Analysis of this new enlarged sample, and also the newly available distribution data of Parapapio from other South African deposits, both tend to support the validity of 3 Parapapio spp., P. jonesi, P. broomi and P. whitei, at Sterkfontein. INTRODUCTION The South African lower Pleistocene limestone deposits have yielded over the years, a very considerable number of fossil cercoplthecoid speCI- mens. In the most recent review of the material recovered (Freedman, 1957), about 1 250 speci­ mens were examined and, since then, several additional significant groups of specimens have been reported (Freedman, 1960, 1961 & 1965). From this material a total of 6 genera, including 14 species, have been described. Of this large and varied cercopithecoid assem­ blage, about 600 specimens come from Sterk­ fontein and almost all of the material has been referred to 3 species of the genus Parapapio. (A few specimens of Cercopithecoides williamsi are present). The genus Parapapio was first described by Jones (1937) for a batch of specimens from Sterkfontein, all of which he included in the type species P. broomi. Subsequently, Broom (1940) decided that the Sterkfontein material included not one but 3 species of Parapapio and he named a slightly larger species P. whitei, and a smaller one, P. jonesi. Material of these 3 species has now also been described from various other South African limestone deposits, namely Swartkrans, Makapan, Taung, Bolt's Farm and Kromdraai (Freedman, 1970). In addition, Broom (1940) re-studied some Papio material from Taung and transferred it to Parapapio (= P. antiquus). Although the various Parapapio species were originally differentiated primarily ~.r.. the basis of dental size variations, subsequent studies utilising more and better material have since revea~~d a number of morpho­ logical cranial and dental differences between the 4 described species. The present study will, however, only be concerned with dental metrical features. Recent work at Sterkfontein (Tobias and Hughes, 1969) has brought to light a further good sample of cercopithecoid specimens. The present paper is concerned with re-examining statistically the specific validity of the 3 Parapapio species at Sterkfontein in terms of the premolar and molar teeth, assessing the newly recovered material and putting on record the premolar and molar size statistics for the total samples now available of each of the 3 Parapapio species from this site. MATERIALS AND ME THODS The Sterkfontein material of Parapapio pre­ viously described by Freedman (1957) was used for the initial analyses of this study. The new rna terial available from Sterkfontein included a total of 565 specimens of which 103 (Appendix 1- 3) were used in the later dental metrical analyses. The area from which the new material came is listed as STW, D13, 1969-1970 (Tobias & Hughes, 1969). This area is a rubble dump near the East Pit and consisted of "large masses of blasted-out breccia" resulting from the early lime working. The rna terial is rich in bone; a considerable variety of mammals and other vertebrates are represented (Tobias & Hughes, 1969). The system of measurements used in this study follows Freedman (1957). Breadth measure­ ments are mesial (bm) and distal (bd); length is maximum (1). Measurements in brackets (Append­ dix) are approximate due to damage. Measure­ ments were taken to the nearest 0,1 mm but mean values are given to 2 decimal places. Where only one decimal place is given in the tables for a 'mean value' only a single observation was available. ANALYSIS A large amount of Parapapio material from Sterkfontein is now available. However, only a small proportion of those specimens can be assigned with certainty to a specific sex and species. Further, because of the fragmentary nature of the specimens, insufficient numbers are available in such gruups for multivariate analyses of, say, 94 P3-M3 or even M1-M3 of either the upper or the lower jaw. Even when analyses of particular single teeth for such comparisons are attempted, most sample sizes are still statistically small. (a) Sexual Dimorphism. Fortunately, one of the features of the genus Parapapio is the very small statistical size differ­ ence between the sexes in the premolars (except P3 length) and the molars. To examine the possibility of pooling the sexes in the comparisons contem­ plated, it was therefore decided to compare the dimensions of these teeth in the two sexes by "t" tests. Where only a single specimen of one sex was available for a particular comparison, a modified "t" test (Simpson, Roe & Lewontin, 1960, p. 183) was used. For this initial study only the previously described and sexed material (Freedman, 1957) was utilised. Tables 1A and 1B show the results of this analysis. Of the 71 "t" tests performed, in the upper dentition only 5 tests were significant at or above the 5% level and in the lower teeth 9 tests revealed this level of significance. Inadequate male samples of P. whitei severely limi ted the number of upper dentition tests possible but the lower teeth comparisons in this taxon confirm the very low degree of dental sexual dimorphism in the genus. It will be noted that, possibly because of the small sample sizes, in 6/14 P. broomi lower dimensions, 4/13 P. jonesi upper dimensions and 11/16 P. whitei measurements, the female mean figure is actually larger than that of the male. This finding is surprising in view of the larger overall size of the male cranium as compared with that of the female in all of the comparisons hitherto possible (Freed­ man, 1957 and 1965). From the results of the analysis of metrical sexual dimorphism in the premolar-molar den­ tition of the 3 Parapapio spp., it would seem that pooling of data from the 2 sexes for inter-specific comparisons can be validated except, as expected, for P31(h). (b) Comparison of the 3 Parapapio spp. at Sterkfontein. Still using the previously described material only, a comparison was next made of the premolar and molar dimensions of the 3 Parapapio spp. from Sterkfontein, using pooled male and female data. The results are shown in Tables 2A and 2B and "t" tests are again used to assess the comparisons. Of the 39 tests on upper tooth dimensions, 31 are significant at the 5% level or higher; using only the molars, 26/27 tests were significant. In the lower tooth comparisons, of the 42 "t" t&sts performed, 31 were significant at the 5% level or better; in the molars, 27/30 tests were significant. The P. whitei- P. broomi comparisons of the lower teeth include the fewest number of significant "t" test results, but even here, 7/10 molar dimensions differ at the 5% level or higher. For each dimension in both the upper and lower teeth the pattern is consistently: P. whitei >P. broomi >P. jonesi. (c) Comparison of the new and old samples of each of the 3 Parapapio spp. from Sterk­ fontein. The 103 newly excavated Parapapio specimens from Sterkfontein used in this study include only a very few fairly complete cranial and mand~bular specimens (Plates 1 and 2); most of the speCImens are only single teeth or small gr.oups of t~e~h. All of these specimens were tentatIvely classIfIed on the assumption of 3 progressi~ely larger sp~cies being present. (In the AppendIx. the~ are hste~ under individual tooth types as P. }OneSl, P. brooml and P. whitei). Assignment was essentially sub­ jective and based on overall size an.d apparent size discontinuities. The 3 groups resultmg (= sample I) were then compared by "t" tests with the ~arlier sample (Freedman, 1957) from the same sI~e (= sample II). Of the 31 upper tooth comparIsons made (Table 3A), . only 3 tests (2 of the~e in l!. whitei) were significant at the 5% level or hIgher; m the 32 lower tooth comparisons (Table 3B), again only 3 significant differences (all in P. jonesi) were found. Because of the paucity of material of P. whitei, only p3, p4, MI and M3 could be compared. Of the 11 tests on these P. whitei teeth, 2 were significant, 1 at the 0,01% level and the other at 0001% level. Comparing the various mean values for 'samples I and II for all 3 species, in the upper teeth the new sample mean values were greater than those of the old sample in all but 7 of the 31 instances; in the lower teeth, the new sample mean was greater than the old in only 11 out of the 32 comparisons made. (d) Statistical summary of the dimensions of the upper and lower pr~molar and molar dim~n­ sions of the Parapapw spp. from Sterkfontem. Tables 4A and B are derived from the combined old and new samples. The numbers of specimens of P. whitei are mostly still small but the P. broomi and P. jonesi samples are now mostly over 10 and half of the samples include over 20 individual measurements. An oddity is the fact that the numbers of P. broomi specimens in the various categories are about double those of P. jonesi in the upper teeth, whilst in the lower tee~h the P. jonesi numbers are mostly greater. It IS not at present clear whether these differences could be due to chance effects or to misassignments. The coefficients of variation (C.V.) and the standard deviations (S.D.) are, for the mo st part, reasonable, except for P3 l(h). These dimensions (and those of the canines) exhibit sexual dimorphism and henc.e high standard deviations and coefficients of van­ ation are to be expected when the 2 sexe~ ~re lumped. The other high valu~s for. these statIs3tIcs are for certain upper tooth dunenslOns (e.g. P b, p4 1 and MIl) of P. whitei. Sample siz.es are small here and the high values result from smgle rather large specimens. (e) Size-frequency distribution of M3 of Para­ papio spp. from Sterkfontein. To investigate further the Parapapio specimens from Sterkfontein, it was decided to re-examine one of the major diagnostic criteria hitherto used for classifying this material. For this purpose the M3 tooth was selected. This tooth was chosen as the hypoconulid makes identification certain, whereas with the other molars there is often some uncertainty as to the position of the tooth in the toothrow, and damaged roots may even make an upper-lower tooth distinction difficult. Further, for this tooth a good sample of over 50 specimens was available. This sample includes the previously described material (Freedman, 1957), which had only been utilized if the sex could be determined, and the new batch of specimens (Appendix) in which every available specimen was measured. Figure 1 is a histogram of M3 1 for 48 specimens of these 2 batches of Parapapio speci­ mens from Sterkfontein. With no selection or subdivision the distribution is obviously bimodal and possibly trimodal. When 2 dimensions of M3 (mesial breadth, bm, and maximum length, 1) are plotted for the 41 available specimens (Fig. 2), there is an apparent cluster below about 10,5 mm breadth and 13,5 mm length, and another less clear group above about 11,0 mm breadth and 16,0 mm length. For making the second subdivision, a cut off at about 16,0 mm appears clear; for the breadth, 11,0 mm seems the logical point on the basis of overall tooth size, as a lower cut off would include teeth very much shorter and hence unreasonably mu ch smaller in overall size. Sub­ divisions at these two length-breadth positions result in 3 groups which correspond with the 3 species postulated for this rna terial. The separation of the 3 species using the breadth and length of M3 becomes more obvious if the 2 dimensions are combined into indices (Table 5). Two indices have been used; the robustness value, bm X 1, and the crown module, (bm + 1)/2 (Goose, 1963). For the robustness values there is only overlap between P. jonesi and P. broomi. This overlap is due to a single large tooth, a male P. jonesi (STS. 258); the next largest tooth in P. jonesi has a robustness value of only 123,69. For the crown module ranges there is no actual overlap but the same single large male P. jonesi tooth just equals the lowest P. broomi figure. DISCUSSION It has been previously noted (Freedman, 1957) that the overall ranges for each dimension (by sex) of the pooled Parapapio material from Sterkfontein (i.e. of all 3 species together) appear to be too great for a single species, when a modern cercopithecoid species (Papio ursinus) is used as a standard. The possibility was also considered that the whole of the Parapapio material might repre­ sent a simple, temporal size increase, as the Sterkfontein rna terial is generally believed to have 95 accumulated over a considerable time. However, during excavation of the site there was no apparent evidence of such a size increase up through the deposit (Robinson- personal communication). Analysis of the distribution of the Parapapio material from the other South African limestone deposits where this genus has been found also tends to discount the possibility that the Sterk­ fontein Parapapio material represents a chrono­ cline. Thus, examination of Table 6 shows that only at Sterkfontein and Makapan are all three Parapapio species present. These are, however, the only 2 sites from which reasonable numbers are available and it is also interesting to note that the 3 species are present at these sites in very similar proportions. (The value of chi square obtained on comparing these two distributions is 1 803 with 2 degrees of freedom, which is well below the 5% level of significance). Although numbers are small at the other sites, it is noteworthy that at Taung only P. jonesi and P. whitei have been recovered, whilst from Bolt's Farm only P. broomi and P. whitei. Of special interest is the fact that only P. jonesi has been found at both Swartkrans and Kromdraai, which are both of more recent age than Sterkfontein and Makapan (Brain, 1958). These latter findings all take on added significance when one recalls that P. broomi is by far the most common form at both Sterkfontein and Makapan, whilst P. whitei is the least common. Maier ( 19 70) has recently describ ed twenty important new specimens from Makapan which he refers to the 5 previously described species from that site. Of this very good material, he refers 6 specimens to P. jonesi, 7 to P. broomi and 2 to P. whitei. He also suggests that, on the basis of this new rna tenal, several prevIOusly described Para­ papio specimens from Makapan require reclassifi­ cation. Maier gives very full anatomical descrip­ tions of the new specimens and is able to extend considerably our morphological knowledge of the populations of these 3 species at that site. However, although detailed measurements of the various specimens are included, he has not attemp­ ted any detailed metrical comparisons. It would seem that a full statistical re-assessment of the 69 Parapapio specimens now known from Makapan (similar to that made here for Sterkfontein) would be extremely valuable. The two populations could then be compared both morphologically and metrically and, hopefully, would resolve a number of the presently contentious aspects. The analyses made earlier, and also the above considerations, all seem to support the probability that the Sterkfontein Parapapio material represents 3 species and the scanty cranial evidence available also tends to support this same hypothesis (Freed­ man, 1957 and 1965). A recent study by Swindler, McCoy and Hornbeck (1966) is also of interest in this connection, as these authors have shown that, on dental metrical data alone, two extant cerco­ pithecoid species, Papio ursinus and Papio anubis, 96 show very considerable overlap for all dental dimensions studied. From the foregoing analysis and review, it would then seem that 3 species of increasing size, P. jonesi, P. broomi and P. whitei, still best account for the Parapapio material hitherto described from Sterkfontein. ACKNOWLEDGEMENTS Professor P. V. Tobias, Department of Ana­ tomy, University of the Witwatersrand, J ohannes­ burg kindly invited one of us (L.F.) to describe this rna terial and the Wenner-Gren Foundation for Anthropological Research, New York made the visit possible by a grant (No. 2402). We should like to thank Dr. W. F. C. Blumer, Department of Anatomy, University of Western Australia, who read the manuscript and made several valuable suggestions. Miss S. Job and Mrs. D. L. Rosman of the Raine Medical Statistics Unit, University of Western Australia assisted with statistical calculations. REFERENCES BRAIN, C. K., (1958). The Transvaal Ape-Man­ Bearing Cave Deposits. Mem. Transv. Mus. 11, 1-13l. BROOM, R., (1940). The South African Pleisto­ cene cercopithecid apes. Ann. Transv. Mus. 20, 89- 100. FREEDMAN, L., (1957). The fossil Cercopithe­ coidea of South Africa. Ann. Transv. Mus. 23, 121- 262. ---, (1960). Some new cercopithecoid speci- mens from Makapansgat, South Africa. Pala­ eont. afro 7,7-45. ----, (1961). New cercopithecoid fossils, in­ cluding a new species, from Taung, Cape Province, South Africa. Ann. S. Afr. Mus. 46, 1-14'. - ---, (1965). Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa. Palaeont. afro 9, 19- 48. ----, (1970). A new check-list of fossil Cerco­ pithecoidea of South Africa. Palaeont. afro 13, 109- 110. GOOSE, D. H., (1963). Dental Measurement: An assessment of its value in anthropological studies. In Brothwell (Ed.): Dental Anthro­ pology. Symp. Soc. Study Hum. BioI. 5, 125- 148 Pergamon Press, Oxford. JONES, T. R., (1937). A new fossil primate from Sterkfontein, Krugersdorp, Transvaal. S. Afr. J. Sci. 33, 709- 728. MAIER, W., (1970). New fossil Cercopithecoidea from the lower Pleistocene cave deposits at Makapansgat limeworks, South Africa. Pala­ eont. afro 13,69- 107. SIMPSON, G. G., ROE, A., AND LEWONTIN, R. c., (1960). Quantitative Zoology. Rev. Ed. Harcourt, Brace & Co., New York. SWINDLER, D. R., McCOY, H. A., AND HORN­ BECK, P. V., (1966). The dentition of the baboon (Papio anubis). In Vagtborg: The Baboon in Medical Research. 2, 133- 150. Univ. of Texas Press, Austin. TOBIAS, P. V., AND HUGHES, A., (1969). The new Witwatersrand University excavation at Sterkfontein. S. Afr. Archaeol. Bull. 24, 158- 169. TABLE 1 A STATISTICAL COMPARISON OF THE MALE AND FEMALE UPPER PREMOLAR AND MOLAR TEETH IN EACH OF THE 3 PARAPAPIO SPP. FROM STERKFONTEIN (in mm) Species Statis- p3 I p4 Ml M2 tics b l(h) b I bm bd I bm bd I bm P. broomi Male 8,15 8,33 8,94 6,63 9,90 9,03 9,95 11,47 10,87 11,38 11,34 Female 7,33 6,49 8,32 6,47 9,52 8,77 9,63 10,86 10,11 10,98 10,53 ** * * *** "t" 4,02 2,54 2,16 1,19 1,86 0,76 1,35 1,62 4,41 1,58 2,01 d.f. 12 13 16 15 14 10 15 9 12 15 9 P. jonesi Male 7,5 4,3 7,7 5,6 9,00 8,60 8,30 10,40 9,60 10,15 10,85 Female 7,05 5,38 7,70 6,03 8,77 7,83 8,87 10,45 8,85 10,13 9,80 "t" 1,15 0,73 0,00 2,71 1,75 1,29 0,88 0,12 1,24 0,03 3,21 * d.f. 3# 2# 2# 3# 2 2 4 1 1 3 3 P. whitei Male - - 8,6 6,7 - - 10,4 - - - - Female 7,87 7,42 8,78 6,90 10,57 10,00 10,56 12,83 11,82 12,72 12,80 "t" - - 0,44 0,23 - - 0,29 - - - - d.f. - - 4# 4# - - 4-it - - - - Ht" test significance levels· * = ° 05 * * = ° 01 * * * = ° 001 # modified "t" test (Sim~son, Roe'& Lew~nti~, 1960, p. 183) M3 bd 1 9,04 11,34 8,77 11,09 0,70 0,76 9 10 8,45 10,25 7,87 9,63 1,29 1,45 3 3 - - 11,35 12,77 - - - - TABLE 1 B STATISTICAL COMPARISON OF THE MALE AND FEMALE LOWER PREMOLAR AND MOLAR TEETH IN EACH OF THE 3 PARAPAP/o SPP. FROM STERKFONTEIN (in mm) Species Statis- P3 P4 MJ M2 tics b l(h) b I bm bd I bm bd I bm P. broomi Male 6,00 14,5 7,15 7,75 7,97 8,03 9,80 10,15 9,93 11,3 3 10,05 Female 5,50 8,37 6,98 7,14 8,27 8,34 9,09 9,93 9,43 11,56 10,13 *** "t'" 1,60 5,60 0,53 1,74 0,79 0,74 1,32 0,41 1,10 0,53 0,17 d.f. 8 8 6 8 6 6 8 7 7 9 5 P. jonesi Male 5,45 14,95 7,01 7,05 8,08 8,08 9,10 9,64 9,54 10,78 9,50 Female 4,90 8,69 6,50 6,66 7,36 7,47 8,42 9,31 9,02 10,39 9,20 ** *** * *** ** * "t" 3,13 9,00 2,10 2,32 5,07 4,25 1,78 1,51 2,38 2,04 1,03 d.f. 12 12 15 13 10 9 13 12 11 13 6 P. whitei Male 5,53 16,4 7,05 7,80 8,30 8,65 9,38 10,65 10,70 12,13 1l,10 Female 5,77 11,33 7,20 7,60 8,67 8,87 10,17 10,63 10,30 12,97 11,00 ** * "t" 1,12 2,40 0,26 0,41 1,04 1,13 5,01 0,08 1,70 2,91 0,67 d.f. 4 2# 3 3 3 3 5 3 3 4 3 - - - -----_.- - - - - "t" test significance levels: * = 0,05, ** = 0,01, *** = 0,001 # modified "t" test (Simpson, Roe & Lewontin, 1960, p. 183) M3 bd bh 9,10 5,95 9,30 6,77 0,44 1,51 8 3 8,67 4,60 8,35 5,04 0,88 1,10 7 4 10,23 6,5 10,40 - 0,39 - 3 - - I 14,83 14,71 0,29 10 13,2 12,82 1,41 7 16,17 16,67 1,34 4 <:J '-I Species P. broomi: Mean P. jonesi: Mean P. whitei: Mean P. broomi "t" vs P. jonesi d.f. P. jonesi "t" vs P. whitei d.f. P. whitei "t" vs P. broomi d.f. TABLE2A STATISTICAL COMPARISON OF THE UPPER PREMOLAR AND MOLAR TEETH (in mm) OF PARAPAPIO SPP. FROM STERKFONTEIN (DATA FROM FREEDMAN, 1957) p3 p4 M' M2 b 1 (h) b 1 bm bd 1 bm bd 1 bm 7,64 7,23 8,59 6,54 9,64 8,81 9,65 11,23 10,49 11,12 10,90 7,10 5,28 7,66 5,90 8,74 8,07 8,62 10,16 9,12 10,14 10,22 7,87 7,42 8,75 6,87 10,57 10,00 10,53 12,83 11,82 12,72 12,80 ** *** *** ** *** ** *** *** 2,06 2,08 2,98 3,92 5,58 3,08 4,78 3,26 5,53 3,60 1,62 19 17 21 20 24 19 28 18 20 20 14 * *** * *** *** *** *** *** *** ** 2,70 1,35 6,21 2,48 9,43 6,21 5,40 8,96 6,95 6,91 4,71 6 6 9 9 9 10 10 9 9 9 5 *** *** ** *** *** *** ** 0,65 0,19 0,52 1,55 4,28 4,60 3,55 5,12 4,87 5,76 3,26 17 17 22 21 21 17 24 19 21 21 11 "t" test significance levels: * = 0,05, ** = 0,01, *** = 0,001 M3 bd 8,89 8,10 11,35 * 2,28 14 *** 7,52 5 *** 4,83 11 1 11,19 9,88 12,77 *** 4,20 15 ** 5,13 6 ** 3,61 13 <.0 00 J: Species P. broomi: Mean P. jonesi: Mean P. whitei: Mean P. broomi "t" vs P. jonesi d.f. P. jonesi "t" vs P. whitei d.f. P. whitei "t" vs P. broomi d.f. TABLE 2 B STATISTICAL COMPARISON OF THE LOWER PREMOLAR AND MOLAR TEETH (in mm) OF PARAPAPIO SPP. FROM STERKFONTEIN (DATA FROM FREEDMAN, 1957) P3 P4 MJ M2 b l(h) b I bm bd I bm bd I bm bd 5,63 11,16 7,06 7,23 8,20 8,24 9,31 10,07 9,72 11,42 10,19 9,30 5,07 10,78 6,77 6,82 7,51 7,59 8,61 9,29 9,02 10,42 9,30 8,40 5,65 12,60 7,14 7,68 8,52 8,78 9,71 10,64 10,46 12,55 11,06 10,30 *** ** *** *** ** *** ** *** *** *** 3,77 0,32 1,68 2,85 4,22 3,88 3,01 3,92 3,10 5,97 4,73 4,95 31 31 31 33 27 26 33 30 29 32 20 24 ** *** *** *** *** *** *** *** *** *** 3,20 1,02 1,44 4,44 4,91 6,07 3,98 5,31 4,88 9,92 11,00 8,17 23 20 23 23 20 18 25 22 21 23 14 15 * * * *** ** *** 0,12 0,74 0,36 1,99 1,33 2,46 1,39 2,20 2,56 4,25 3,56 4,14 18 17 16 18 15' 16 20 16 16 19 14 17 "t" test significance levels: * = 0,05, ** = 0,01, *** = 0,001 M3 bh 6,22 4,95 6,50 *** 4,96 15 *** 5,06 8 0,58 9 I 14,72 12,71 16,42 *** 8,31 27 *** 12,01 17 *** 5,96 20 \.0 \.0 Species P. jonesi P. broomi P. whitei TABLE 3 A COMPARISON OF THE UPPER PREMOLAR AND MOLAR TEETH (in mm) OF TWO SAMPLES OF 3 SPECIES OF PARAPAPIO FROM STERKFONTEIN (I = NEW SAMPLE; II = 1957 SAMPLE) Statis- p3 p4 Ml M2 tics b l(h) b 1 bm bd 1 bm bd 1 bm M (I) - - 7,87 5,95 8,55 8,10 8,93 10,44 9,26 10,50 9,85 M(II) 7,10 5,28 7,77 6,11 8,74 8,07 8,62 10,16 9,12 10,13 10,08 "t" - - 0,67 0,4~ 0,64 0,06 0,73 1,05 0,62 1,61 0,71 d.f. - - 8 8 7 7 11 8 8 12 10 M(I) . 7,70 7,80 8,45 6,75 9,82 8,92 9,95 11,43 10,50 11,21 10,60 M(Il) 7,65 7,16 8,60 6,54 9,64 8,81 9,65 11,23 10,49 11,18 10,98 "t" 0,21 0,85 0,55 1,19 1,00 0,47 1,19 0,62 0,05 0,15 0,98 d.f. 20 19 24 23 23 18 24 19 22 26 19 M(I) 9,5 8,4 9,93 7,90 11,1 10,7 13,2 - - - - M(Il) 7,87 7,42 8,75 6,87 10,57 10,08 10,53 12,83 11,82 12,72 12,80 *** ** "t" 3,74 0,39 5,44 1,67 1,16 1,07 5,43 - - - - d.f. 2 3 7 6 3 4 5 - - - - "t" test significance levels: * = 0,05, ** = 0,01, *** = 0,001 M3 bd 8,86 8,07 * 2,99 9 8,92 9,09 0,45 18 - 11,35 - - 1 10,00 9,85 0,70 11 10,87 11,21 1,16 20 - 12,72 - - ..... o o Species Statis- tics P. jonesi M(I) M(ll) "t" d.f. P. broomi M(I) M(ll) "t" d.f. P. whitei M(I) M(ll) "t" d.f. TABLE 3 B COMPARISON OF THE LOWER PREMOLAR AND MOLAR TEETH (in mm) OF TWO SAMPLES OF 3 SPECIES OF PARAPAPIO FROM STERKFONTEIN (I = NEW SAMPLE; II = 1957 SAMPLE) P3 P4 MJ M2 b I(h) b I bm bd I bm bd I bm bd 5,15 14,55 6,17 6,97 7,23 7,40 9,18 8,70 8,28 10,44 8,99 7,92 5,07 10,78 6,69 6,82 7,51 7,59 8,61 9,29 9,02 10,42 9,30 8,40 * ** * 0,25 1,58 1,76 0,76 1,09 0,73 1,62 2,72 3,31 0,11 1,91 2,35 19 18 24 22 18 16 23 26 25 27 16 18 4,8 15,0 7,0 7,1 7,97 8,03 9,87 9,20 9,00 11,40 9,82 8,96 5,63 11,16 7,06 7,23 8,20 8,24 9,31 10,00 9,72 11,42 10,19 9,30 1,92 1,04 0,15 0,27 0,78 0,72 1,28 2,09 1,63 0,05 1,67 1,75 13 14 12 14 13 14 16 13 13 15 21 24 - - - - - - - - - - 11,4 10,4 5,65 12,60 7,14 7,68 8,52 8,78 9,71 10,64 10,46 12,55 11,06 10,30 - - - - - - - - - - 2,05 0,22 - - - - - - - - - - 4 4 "t" test significance levels: * = 0,05; ** = 0,01, *** = 0,001 M3 bh 5,00 4,95 0,26 13 6,22 6,22 0,02 18 6,1 6,50 0,77 1 I 12,75 12,71 0,17 19 14,52 14,72 0,92 26 15,3 16,42 2,10 5 ..... o ..... Species Statis- tics P. jonesi M N S.D. C.V. P. broomi M N S.D. c.v. P. whitei M N S.D. C.V. TABLE 4 A STATISTICAL SUMMARY OF THE DIMENSIONS OF THE UPPER PREMOLAR AND MOLAR TEETH (in mm) OF PARAPAPIO SPP. FROM STERKFONTEIN p3 p4 Ml M2 b l(h) b 1 bm bd 1 bm bd 1 bm 7,10 5,28 7,80 6,08 8,70 8,08 8,69 10,30 9,19 10,29 9,97 5 4 10 10 9 9 13 10 10 14 12 0,31 1,63 0,20 0,46 0,36 0,55 0,64 0,42 0,34 0,46 0,55 4,4 30,9 2,6 7,6 4,1 6,8 7,3 4,1 3,8 4,4 5,5 7,66 7,31 8,57 6,59 9,68 8,84 9,72 11,29 10,49 11,19 10,87 22 21 26 25 25 20 26 21 24 28 21 0,48 1,46 0,58 0,38 0,37 0,43 0,54 0,68 0,51 0,56 0,80 6,2 20,0 6,7 5,8 3,9 4,9 5,5 6,0 4,8 5,0 7,4 8,28 7,62 9,14 7,13 10,68 10,18 10,91 12,83 11,82 12,72 12,80 4 5 9 8 5 6 7 6 6 6 2 0,87 1,97 0,66 0,85 0,42 0,54 1,09 0,55 0,77 0,65 - 10,5 25,9 7,2 11,9 4,0 5,2 10,0 4,3 6,5 5,1 - M3 bd 8,43 11 0,58 6,9 9,05 20 0,70 7,7 11,35 2 - - 1 9,93 13 0,38 3,8 11,12 22 0,61 5,5 12,77 3 - - - - ...... o I\;) Species P. jonesi P. broomi P. whitei TABLE 4 B STATISTICAL SUMMARY OF THE DIMENSIONS OF THE LOWER PREMOLAR AND MOLAR TEETH (in mm) OF PARAPAPIO SPP. FROM STERKFONTEIN Statis- P~ P4 Ml M2 tics b l(h) b 1 bm bd 1 bm bd 1 bm M 5,08 11,16 6,61 6,85 7,47 7,56 8,72 9,10 8,77 10,43 9,18 N 21 20 26 24 20 18 25 28 27 29 18 S.D. 0,40 3,33 0,56 0,38 0,40 0,41 0,73 0,60 0,65 0,42 0,36 C.V. 7,8 29,8 8,5 5,6 5,4 5,4 8,3 6,5 7,4 4,1 4,0 M 5,57 11,40 7,06 7,22 8,15 8,20 9,40 9,95 9,62 11,42 10,00 N 15 16 14 16 15 16 18 15 15 17 23 S.D. 0,46 3,59 0,39 0,44 0,46 0,44 0,70 0,60 0,62 0,52 . 0,55 C.V. 8,2 31,5 5,5 6,2 5,6 5,4 7,4 6,1 6,4 4,6 5,5 M 5,65 12,60 7,14 7,68 8,52 8,78 9,71 10,64 10,46 12,55 11,12 N 6 4 5 5 5 5 7 5 5 6 6 S.D. 0,23 2,80 0,45 0,37 0,39 0,22 0,46 0,21 0,31 0,55 0,19 C.V. 4,1 22,2 6,3 4,8 4,6 2,5 4,7 2,0 3,0 4,4 . 1,7 M3 bd bh 8,21 4,97 20 15 0,49 0,35 6,0 7,1 9,14 6,22 26 20 0,51 0,53 5,6 8,5 10,32 6,37 6 3 0,38 0,38 3,6 5,9 1 12,73 21 0,54 4,3 14,63 28 0,57 3,9 16,26 7 0,62 3,8 ..... o vo 104 TABLE 5 OBSERVED RANGES FOR TWO INDICES (ROBUSTNESS VALUE & CROWN MODULE) FOR 41 M3 TEETH OF THE 3 PARAPAPIO SPP. FROM STERKFONTEIN Index P. jonesi P. broomi P. whitei (N = 15) (N _= 20) (N = 6) Robustness value 103,32 - 130,00 126,00 - 166,92 172,05 - 184,80 Crown module Species P. jonesi. , . P. broomi P. whitei 10,35 - 11,50 11,50 - 13,15 13,30 - TABLE 6 NUMBER OF SPECIMENS OF PARAPAPIO DESCRIBED FROM VARIOUS SOUTH AFRICAN LIMESTONE DEPOSITS Taung Sterkfontein Makapan* Swartkrans Kromdraai (lim ew or ks ) (faunal site) 1 25 15 3 2 0 43 34 0 0 3 14 5 0 0 * Excluding Maier (1970) 13,85 Bolt's Farm 0 5 2 (a) Upper teeth Spec. p3 No. b STW 18 8,1 \ Spe.:. p4 No. b r S1W 18 - 23 7,8 27 8,0 77 7,8 (b) Lower teeth Spec. P3 No. b STW 3 5,2 57 5,1 Spec. p4 No. b STW 3 6,2 9 6,2 46 6,3 78 6,0 APPENDIX 1 MEASUREMENTS (in mm) OF NEW PARAPAPIO MATERIAL FROM STERKFONTEIN 1. SPECIMENS TENTATIVELY ASSIGNED TO P. JONES! Spec. M' Spec. M2 Spec. l(h) No. bm bd 1 No. bm bd 1 No. bm . 7,5 STW 18 - - 9,3 STW18 11,1 9,6 11,0 STW 18 10,5 23 9,1 8,6 9,6 20 10,2 9,4 10,7 20 9,7 53 8,0 7,6 7,9 23 10,3 9,0 10,6 24 - 24 - - 10,2 27 9,9 l(h) 27 10,6 9,3 10,4 49 10,0 34 10,0 9,0 10,1 50 10,1 7,0 72 9,4 81 9,4 - 6,1 5,8 -- ---- --_. --_ .. _- - -- Spec. M, Spec. M2 Spec. l(h) No. bm bd 1 No. bm bd 1 No. bm 15,4 STW 3 (7,6) (7,9) 10,0 STW 3 8,8 8,3 11,2 STW 3 8,8 13,7 9 - - 8,4 9 8,9 7,9 10,1 9 8,9 22 (7,0) (7,2) 8,3 11 (9,3) 8,8 10,4 11 - 46 - - 9,2 15 8,8 8,6 11,0 22 9,0 82 7,1 7,1 1.0,0 22 8,6 8,0 10,1 33 9,1 l(h) 33 9,1 - 10,3 43 9,2 46 - 8,5 9,8 53 9,5 51 7,8 8,0 10,2 56 8,4 7,2 64 9,0 8,4 10,8 6,5 65 8,0 8,0 10,5 7,1 7,1 M3 bd - 8,2 - 8,2 9,0 9,0 9,0 9,1 -- - M3 bd 7,5 7,6 8,4 8,1 7,7 8,1 8,6 7,4 1 10,4 10;0 9,8 10,1 10,3 9,8 9,6 10,1 bh 4,8 4,6 5,0 - 4,6 5,4 5,2 5,4 1 13,0 12,5 12,8 12,6 13,0 13,0 12,6 12,3 ...... o (,Jl APPENDIX 2 MEASUREMENTS (in mm) OF NEW PARAPAPIO MATERIAL FROM STERKFONTEIN (continued) 2. SPECIMENS TENTATIVELY ASSIGNED TO P. BROOM! (a) Upper teeth Spec. p3 Spec. Ml Spec. M2 Spec. No. b l(h) No. bm bd I No. bm bd I No. bm STW 10 7,2 7,4 STW 2 10,1 9,1 10,3 STW 2 11,9 11,2 12,1 STW 12 (11,5) 14 7,7 5,9 6 9,6 8,5 10,3 10 11,4 10,3 11,4 58 10,8 58 7,6 9,0 10 9,7 9,1 10,5 12 11,0 10,1 10,5 30 9,3 84 8,2 8,6 58 9,5 9,0 9,5 14 11,2 9,9 11,1 73 11,0 67 10,0 8,8 9,5 25 - 10,4 11,2 87 11,1 74 10,1 9,0 9,6 58 12,1 11,2 11,7 63 10,5 10,2 10,5 67 12,4 . 11 ,4 12,1 Spec. p4 No. b l(h) STW 10 8,5 7,2 12 8,7 5,9 14 8,6 7,5 58 8,5 6,6 84 7,9 6,1 I 2 8,5 7,2 M3 bd 9,0 9,0 - 9,4 9,0 I 10,7 11,2 11,7 10,7 10,8 ...... o O"l APPENDIX 3 MEASUREMENTS (in mm) OF NEW PARAPAPIO MATERIAL FROM STERKFONTEIN (continued) 2. SPECIMENS TENTATIVELY ASSIGNED TO P. BROOMI (continued) (b) Lower teeth Spec. P3 Spec. M, Spec. M3 Spec. No. b I(h) No. bm bd I No. bm bd bh I No. STW 70 4,8 15,0 STW 61 8,3 8,3 10,4 STW 1 9,8 8,7 5,8 14,1 STW 41 83 7,5 7,7 9,2 8 9,6 8,6 5,9 14,8 42 1 (8,1) (8,1) 10,0 13 11,0 9,4 - 14,7 47 Spec. P4 17 9,5 8,2 6,5 13,8 83 No. b I(h) 32 10,4 8,8 6,0 14,2 Spec. M2 37 10,1 9,7 6,0 14,5 38 10,2 9,8 7,0 14,6 STW 1 7,0 7,1 No. bm bd I 40 9,3 8,3 6,0 14,7 STW 1 9,5 9,0 11,0 69 8,9 8,9 11,7 83 9,5 9,1 11,1 3. SPECIMENS TENTATIVELY ASSIGNED TOP. WHITEI (a) Upper teeth Spec. p3 Spec. p4 Spec. M' M2 No. b I(h) No. b I(h) No. bm bd I Nil STW 21 9,5 8,4 STW 31 10,1 - STW 21 (11,1 ) (10,7) 13,2 35 10,0 8,6 M3 21 9,7 7,2 Nil - - - - ---- (b) Lower teeth P3 M, Spec .. M3 No. bm bd bh I Nil Nil STW 36 11,4 10,4 6,1 15 ,3 P4 M2 Nil Nil bm bd 9,4 9,0 9,9 9,4 9,5 8,7 9,2 8,9 M3 (contd.) bh 5,9 6,8 6,8 5,7 I 14,6 15,6 14,8 13,8 ...... o " 108 V'I C Q) E u Q) (L V'I "- 0 ~ Q) ...a E ::> c 12- 11 - 10 - g- a- 7 - 6- 5 - 4 3 2 1 12-1 12-6 13·1 13·6 14·1 length 14·6 15·1 15·6 16·1 Fig. 1. m.m. Histogram of the length (1) of 48 specimens of M3 of Parapapio spp. from Sterkfontein. 16·6 17·1 12· 0 11·0 E E c: .- 10·0 E ~ L -." 0 9·0 Q) L- ~ 8·0 0 0 0 0 0 0 0 0 0 0 0 0 12·0 13·0 x 0 0 0 x x x x x 14·0 length (I) 0 0 x x ~ x x x x x x x x X x 15·0 16·0 In m.m. Fig. 2. Scattergram of the length (1) and mesial breadth (bm) of 41 specimens of M3 of Parapapio spp. from Sterkfontein. O - P. j onesi; X - Po broomi; O - P. whitei. 109 0 61 0 17·0 110 Plate 1 From above downwards, dorsal, lateral and ventral views of P. broomi female cranium (STW 12). I "MI. 0 .. i •• I Plate 2 Top: Palates of P. broomi females (left, STW 58; right, STW 10) Centre: Lateral and occlusal views of P. jonesi male mandible (STW 3). Bottom: Lateral and occlusal views of P. broomi? female mandible (STW 1). III