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Home , Gigantopithecus, Meganthropus

Gigantopithecus and Its Relationship to

DAVID W. FRAYER Department of Anthropology, University of Michigan, Ann Arbor, Michigan 481 04

KEY WORDS . Australopithecus . P. . Hominid origins.

ABSTRACT Gigantopithecus blacki and G. bilaspurensis are compared to P. gorilla and Australopithecus. The total morphological pattern of Giganto- pithecus is more similar to Australopithecus than to P. gorilla. Two major points are raised. (1) G. blacki might be considered an aberrant hominid rather than an aberrant pongid. (2) G. bilaspurensis can be considered an equal- ly likely candidate, along with Ramapithecus, for possible hominid ancestry.

Numerous taxonomic and evolutionary topithecus and Australopithecus. Further- statements have been made concerning more, a direct comparison between the the Gigantopithecus. These inter- Middle Pliocene G. bilaspurensis and Aus- pretations have portrayed Gigantopithecus tralopithecus has not been done, although as ancestral to hominids, as an over-spe- Simons and Chopra (‘69a) allude to some cialized side-branch of hominids, or as basic similarities. The metric data pre- an aberrant pongid, unrelated to hominid sented below combined with a review of evolution. Weidenreich was the main pro- morphological data are intended to clarify ponent of the ancestor/descendent rela- these relationships. tionship between Gigantopithecus and DATA “” ( = Australopithecus) (‘45, ’46, ’49). More recently, Eckhardt (‘71, Dental and mandibular measurements ’72) has reopened the possibility of Gigan- of P. gorilla, Australopithecus, and Gigan- topithecus as a hominid forbear. Woo (‘62) topithecus appear in Appendices 1-5. and von Koenigswald (‘52, ’58) suggested Gorilla tooth dimensions were taken by that Gigantopithecus should be considered Paul E. Mahler and Milford H. Wolpoff. a hominid, but that the genus represents All mandibular measurements were taken an over-specialized side-branch which did by the author. Since all the Cleveland not contribute to evolu- Museum of Natural History gorilla speci- tion, but became extinct. Both of these mens were wild-shot, reliable sex deter- schemes consider only G. blachi and not minations are possible. The sex of the the recently discovered Middle Pliocene specimens listed in Appendices 4-5, then, G. bilaspurensis. being determined in the field from the Simons and Pilbeam (‘65) reviewed the carcass, is quite accurate. G. blachi material and concluded that the Except for noted cases, all australopith- genus represented an aberrant pongid ecine measurements were taken on the with peculiar hominid parallelisms. Si- original specimens by Milford H. Wolpoff. mons’ subsequent discovery of the Indian These are classified into gracile Gigantopithecus (G. bilaspurensis) further and robust categories based either on the confirmed, they felt, this aberrant pongid specific site (e.g., Sterkfontein = gracile) status (Pilbeam, ’70; Simons and Ettel, or on published accounts (e.g., robust and ’70; Simons and Chopra, ’69a,b; Simons gracile forms from Omo). The gracile and Pilbeam, ’72). sample includes South African specimens Throughout this debate, however, little generally described as A. africanus and substantive information has been given 1 This paper is a revised version of paper delivered which would indicate the metrical and at the 1971 meeting of the American Association of morphological similarity between Gigan- Physical Anthropologists,

AM, J. PHYS. ANTHROP.,39: 413-426 413 414 DAVID W. FRAYER East African Lower specimens mates of posterior tooth size in described as . ER-992 has been in- are more reliable when considering only cluded in the gracile sample with reserva- the summed areas of P4-M3. I have for- tions. The robust sample consists of speci- mulated the same indices using summed mens published as A. “robustus” and A. P3-Ms length and areas and have obtained “boisei.” Severely crushed and distorted results similar to those presented in ta- specimens are not included in these groups, ble 1. The inclusion of all the measure- but in some cases teeth with post-mortem, ments provides data for analysis in either matrix-filled cracks are utilized. In these manner. specimens the dimensions of the crack Each index in table 1 intends to show have been subtracted from the overall morphological characteristics through dimension, rendering a good representa- metric evaluation. The three genera are tion of the true tooth size. The notable represented by measurements presented case in this regard is the East African in indices allowing direct comparison of robust specimen, ER-818. Formal descrip- metric/morphological relationships. tions of these newly discovered East Afri- Bicanine breadth divided by the length can hominids appear in Leakey (’71, ’72) of the cheekteeth (P4-Mz) compares the and Leakey et al. (‘71, ’72). maximum external breadth outside the With the exception of bicanine breadth, canines to the length of the posterior all measurements for Gigantopithecus tooth row. The size of the canine (length blachi are taken from Woo (‘62). To my X breadth) is normalized against the knowledge published mandibular dimen- summed areas of the posterior teeth, sions for G. bilaspurensis do not exist. while the area of the lower second Consequently, all dimensions on the Bilas- is compared to the area of the lower first pur were taken on the Yale- . I have also compared lower Is area Peabody Museum cast. Checking dental with the summed posterior tooth areas dimensions from the cast with those pub- as well as lower Mz area. All indices fol- lished by Simons and Chopra (’69a), the low the same pattern, but for the sake of Yale-Peabody Museum case is within 4% brevity, have not been included in this of the dimensions on the original speci- paper. Maximum symphyseal length is men. Mandibular dimensions taken from considered in regard to the height of the the cast for the Bilaspur mandible are mandible at Mn. This index is somewhat assumed to be within 4% of the true different from that described in Simons values. Bicanine breadth on G. blachi is and Chopra (‘69a), but again demonstrates similarly accurate. Finally, dental dimen- the same morphological relationship as sions for all Gigantopithecus mandibles their index of maximum symphyseal length are published by Simons and Chopra (‘69a) divided by mandible height multiplied by and are not duplicated here. breadth at Mz and by the summed pos- METHOD terior cheek teeth area. Maximum sym- physeal length divided by corpus height at The indices presented in table 1 are in Mz better separates Australopithecus from most cases those used by Simons and P. gorilla and, for this reason, is used in Chopra (‘69a) in their taxonomic discus- this analysis. The summed areas of P4-Mz sion of G. bilaspurensis and G. blachi. In divided by height X breadth of the man- place of P3-M3 length and P3-M3 summed dible at Mz normalizes the cheek teeth areas, I have chosen to substitute P4-M2 relative to the robusticity of the mandible length and area. This substitution allows under Mz. Area (1 X b) of MI divided by for a larger sample size in Gigantopithecus height X breadth of the corpus under Mz (G. blachi I and I1 lack lower third molars) provides a further check. and in Australopithecus. It also eliminates These indices are used to demonstrate the comparison of lower third morphological patterns through metric between known pongids and hominids. analysis. They provide excellent sorting Since the lower third in is criteria for separating a mesiodistally elongated tooth set at an from gorillas. Student’s t-tests performed angle to the tooth row and is not directly on these two genera (Australopithecus involved in grinding and crushing, esti- and P. gorilla) using the six indices all GIGANTOPITHECUS AND AUSTRALOPITHECUS 415 showed significant differences at the 0.01 contra these authors, does not specifically level. It follows, then, since the indices indicate that gigantopithecines are “aber- separate australopithecines from gorillas, rant apes.” Rather, if one has to resort that they are useful criteria for analyzing to such terminology, the close metric sim- other hominoid, and possibly related ilarity between G. blacki, G. bilaspurensis genera. and Australopithecus would better de- RESULTS scribe an aberrant hominid. Three major points can be readily seen (3) The third observation apparent from from an examination of table 1. (1) Aus- table 1 is that G. blacki and G. bilaspuren- tralopithecus differs widely from gorillas, sis show some different metric relation- a fact which has been discussed above and ships. The relative bicanine breadth and by numerous authors beginning with relative symphyseal length are both small- Dart (‘26). Different cultural and dietary er in G. bilaspurensis than in G. blacki I, adaptations are likely the major reasons I1 or 111, and one would expect the index for this metric difference. The morpho- describing lower incisor area to differ logical specialization of Australopithecus also. In some cases G. bilaspurensis closely to small object feeding has been consid- resembles Australopithecus. ered in detail by Jolly (‘70). The infie- This similarity is further emphasized quency of range overlap between Australo- when comparing G. bilaspurensis with two pithecus and P. gorilla provides new robust australopithecines from East Africa evidence for determining the phylogenetic (see table 2). Here the major differences positions of G. blacki and G. bilaspurensis. between the Bilaspur mandible, Omo 7, (2) Except in two cases (relative sym- and ER-729 relate to absolute canine size. physeal length and relative p4-M~ area, The smaller size of the bicanine breadth both in G. blacki 11), indices describing in G. bilaspurensis reflects the size of the Gigantopithecus mandibles and tooth size canines rather than the size of the . fall completely within the range of Aus- The internal bicanine breadth (measured tralopithecus. Both G. blacki and G. bi- from lingual surface on the canine) dem- laspurensis overlap extensively with Aus- onstrates this clearly. The size of the tralopithecus but seldom with P. gorilla. mandibular corpus under Mn is somewhat In those cases where Gigantopithecus more massive in G. bilaspurensis than in indices have values within the P. gorilla robust australopithecines although recent range, Australopithecus also has values East Rudolf finds almost equal it in cross- which overlap with P. gorilla. sectional area, or are larger (see Appen- It is interesting to note that the largest dix 1). At least two have mandibles greater mandible of Gigantopithecus (G. blacki 111), in breadth. No gorilla specimen even ap- an assumed male (Simons and Chopra, proaches G. bilaspurensis or these large, ’69a; Woo, ’62), is well below the range robust australopithecines in mandibular in all indices describing male gorillas. In corpus cross-sectional area at Mz. canine size, G. blacki I11 has canines Measurements of the mandible and den- which are relatively smaller than the ca- tition of G. bilaspurensis and Australo- nines of the presumed female, G. blacki I. pithecus, then, closely resemble each in canine size of G. other. Both have absolutely and relatively blacki does not follow the common pongid narrow bicanine breadths bounded by condition where little or no overlap in relatively small canines. The lower incisors relative canine size is characteristic. Con- are completely unlike gorillas, being con- sequently, relative canine size cannot be siderably reduced. The maximum sym- used to discriminate male from female physeal length is greater than the corpus Gigantopithecus. height at the lower second molar, but not The general similarity of Gigantopithe- to the extent of the condition seen in cus to Australopithecus and its dissimi- gorillas. The posterior teeth are both rela- larity in all indices, as well as sexual tively and absolutely large as is the cross- dimorphism based on relative canine size, sectional area of the mandible under Mn. to gorillas suggests at least a dietary sim- The major metrical difference between the ilarity to Australopithecus (Simons and purported aberrant pongid and hominids Ettel, ’70; Pilbeam, ’70; Jolly, ’70), but, is the size of the lower canine. TABLE I

Summary indices for Gigantopithecus, Austrdopithecus, and gorilla. Sample size is enclosed in brackets. Standard deviations (s.d.) are included where appropriate. All indices are multiplied by 100

Bicanine Br. Canine area Area 12 Max. symph. It. Area P4-M2 Area MI Length PI-Mz Area P4-Mz Area MI Ht. at Mz Ht. X br. at Mz Ht. X br. at Mz G. blacki (I) 89.7 20.9 24.7 126.7 47.1 16.4 G. blacki (11) - 20.8 145.3 63.2 21 .o G. blacki (111) 17.9 12.7 99.3 22.9 134.0 40.0 G. bilaspurensis (IV) 20.2 13.5 Australopithecus (gracile) 73.2 119.8 41.4 Mean 17.8 (8) 99.0 (2) 19.5 (5) = 3.6 s.d. =2.0 30.7 (7) 114.3= (3)12.5 56.6= (4) 8.1 s.d. =2.7 Range s.d. (14.G21.3) s.d. =4.1 (100.6-125.0) (45.w34.4)s.d. (1 5.1-22.2) Australopithecus (robust) (98.7-99.4) (23.3-37.1) s.d. Mean 79.6 (7) 12.2 (8) 123.2 (5) 56.0 (8) 18.8 (8) s.d. = 12.1 s.d. = 2.9 21.4= (4) 3.7 s.d. = 14.1 s.d. = 9.7 s.d. = 2.9 Range (58.2-95.2) (8.3-16.6) (16.0-24.1)s.d. (107.9-1 38.2) (42.4-71.9) ( 13.6-2 1.7) Australopithecus (combined) Mean 84.8 (9) 15.0 (16) 27.3 (11) 119.9 (8) 56.2 (12) 19.1 (13) s.d. = 13.1 s.d. = 3.8 s.d. =6.0 s.d. = 13.4 s.d. = 8.8 s.d. =2.7 (58.2-99.4) (8.3-21.3) (1 6.0-37.1) (100.61 38.2) (42.4-71.9) (1 3.6-22.2) P. gorilla (male) Range 129.9 (20) 42.3 (20) 44.5 (18) 179.6 (20) 79.8 (20) 26.6 (20) Mean s.d. = 10.7 s.d. = 4.9 s.d. = 5.7 s.d. = 13.6 s.d. = 11.1 s.d. = 3.8 (1 13.5-154.8) (31.8-53.0) (35.7-54.6) (156.6-213.8) (62.6-105.2) (20.3-35.1) P. gorilla (female) Range 112.7 (20) 23.9 (39) 41.9 (19) 164.0 (20) 82.4 (19) 28.3 (20) s.d. = 9.2 s.d. =2.2 s.d. = 5.3 s.d. = 11.7 s.d. = 17.6 s.d. = 5.8 Mean (96.0-129.8) (20.8-29.5) (31.6-49.7) (142.1-186.3) (62.1-1 13.5) (18.4-39.1) P. gorilla (male and female) Range 121.3 (40) 33.4 (39) 43.1 (37) 171.8 (40) 81.1 (39) 27.4 (40) s.d. = 13.2 s.d. = 1 .O s.d. = 5.6 s.d. = 14.8 s.d. = 14.1 s.d. =4.9 Mean (96.0-154.8) (20.s53.0) (3 1.6-54.6) (142.1-213.8) (62.1-113.5) ( 1 9.9-39.1 ) Range GIGANTOPITHECUS AND AUSTRALOPITHECUS 417

TABLE 2 Tooth measurements and indices for two East %frican australopithecines and G. bilaspurensis Omo 7 KNM-ER-729 G. bilaspurensis Areas of mandibular teeth Canine 74.9 85.9 148.8 PM3 182.0 150.5 172.3 PM4 221.1 205.9 190.0 M1 314.2 244.9 241.4 ..M2 291.6 342.0 305.6 M3 269.4 383.4 315.5 Summed cheek teeth 1278.3 1326.7 1224.8 Area P4-M2 loo Ht. X br. at M2 53.8 58.8 41.4 Bicaninebr. 58.2 1 76.0 73.2 2 Length P4-M2

Max. symph. It. 107.9 1 112.7 119.8 2 Ht. at M2 Minimum bicanine breadth 20.2 22.5 21.0 *

I Measurement from cast belonging to F. C. Howell. 2 Measurement from Yale-Peabody cast.

Morphological details of Australopithe- depth posteriorly. This characteristic is cus and G. bilaspurensis are also quite similar to East African robust forms, similar. Besides the metrical details of ER-403, 404, and 726, but different from the horizontal ramus under M2, other others. ER-818, for example, sharply de- characters of the mandibular body can creases posteriorly in horizontal ramus be duplicated in australopithecines. Sym- depth. physeal cross-sections of Natron, ER-729, The two major differences between G. and ER-818, as well as other early hom- bilaspurensis and Australopithecus in inids, show essentially the same morpho- mandibular morphological details are the logical features. The middle symphyseal angle of the symphyseal region and the area is buttressed by two transverse tori, shape of the dental arcade. In most aus- one running superior and one inferior tralopithecines the symphyseal angle is to the genial-glossal fossa. Above the su- nearly perpendicular to the horizontal perior torus is a shelf-like area which is ramus, while in the Bilaspur mandible characteristic of most australopithecines the area is at a 45" angle to the horizontal and pongids, both and recent. The ramus. It is noteworthy that Ramapithecus inferior transverse torus of G. bilaspuren- also has a non-perpendicular symphyseal sis forms a short shelf, more ver- angle (Andrews, '71). The contour of the tical than in P. gorilla with a relatively tooth row in the Bilaspur mandible is V- higher placement of the genial-glossal shaped, resembling its probable mid-Mio- fossa. cene dryopithecine ancestors which all The origin of the ascending ramus in have anteriorly convergent dental arcades. G. bilaspurensis arises at MI a consider- This feature differs from Australopithecus able distance lateral to the buccal aspect and is not unlike the palatal shape in of the tooth. The result of the condition Ramapithecus. With decrease in size of is a wide buccinator groove similar to the canines and shortening of the mandi- those found in some robust australopithe- ble the palatal contour of G. bilaspurensis cines. In the later G. blachi mandibles, would probably resemble Australopithecus. the ascending ramus begins further pos- G. bilaspurensis has a P4, MI, Mz, and terior at Mz and the buccinator groove is M3 of similar shape to Australopithecus, narrower and less marked. The horizontal but the teeth lack crenulations on the oc- ramus of G. bilaspurensis increases in clusal surfaces. Unworn molars are longer 418 DAVID W. FRAYER than broad with bulging lingual and buc- Chopra, ’69a). A steeply inclined wear cal aspects (see table 3). Comparing heav- facet on the lower canine would indicate ily-worn molars of Australopithecus to that a maxillary canine, through occlu- those of G. bilaspurensis as Simons does sion with the mandibular canine, was act- (‘72:258) only confuses the similarity of ing in life to sharpen the distal edge of molar shape with australopithecines. On the lower canine. Since the lower canines the average, G. bilaspurensis has length/ of G. bilaspurensis are truncated more breadth ratios at the upper end of the than sharpened and since the lower third Australopithecus range or just slightly premolar lacks a large well-defined mesial beyond it. contact facet, it is reasonable to infer that Occlusally, the molars of G. bilaspuren- the maxillary canine was not large, nor sis are less similar to Australopithecus, similar in function compared to gorillas. but are also completely unlike P. gorilla. Rather, since the mandibular canine was The occlusal surfaces of the teeth in the truncated during life, masticatory actions Bilaspur mandible are characterized by performed at the canine appear to be low, broad cusps with shallow fovea sep- more involved with grinding, than with arating each . These teeth are not shearing and gripping functions. The ab- characterized by highly crenulated sur- solute size, wear, and morphology of the faces, although the cusp pattern can be mandibular canines on the Bilaspur man- duplicated in several robust australo- dible, then, do not completely match either pithecines. P. gorilla or Australopithecus. The occlusal morphology on the lower (2) As mentioned above, the third lower molars is strikingly similar to the descrip- premolar of G. bilaspurensis also differs tion of the Ngorora M2 found in Pliocene from hominids. This tooth in addition to deposits in East Africa (Bishop and Chap- having only a very small mesial contact man, ’70). This upper molar described by facet, is set at an angle to the tooth row Leakey is also bunodont with low rounded and displays at least three cusps. Most cusps and distinct fovea separating the investigators have argued that this tooth cusps, and appears to be about the same is pongid-like, considering it somewhat geological age as G. bilaspurensis. Due to modified from a true sectorial lower third the lack of other African middle Pliocene premolar. It is important to note, how- forms, this relationship is certainly ten- ever, that the model of a Pliocene hominid uous. ancestor with bicuspid lower third pre- Another similarity in dental morphology molars, is no longer supported by the data, is the “plate-like” premolar and molar since the Fort Ternan ramapithecine has roots of both genera. From breaks and a sectorial lower third premolar (Andrews, erosions in the Bilaspur mandible one can ’71). This tooth is also set at an angle to observe a left Pq anterior root, a left MI the tooth row, and further differs from anterior root, and a right M:3 posterior G. bilaspurensis in having a long, mesial root, all of which are “plate-like.’’ Leakey contact facet for occlusion with the max- et al. (‘71) have noted “plate-like” roots illary canine. The morphology of P3 on in ER-404, and Wolpoff has observed them G. bilaspurensis can not be used to ex- in the originals of ER-725, 726, and 733a, clude it from the status of a possible hom- and ER-818 (personal comm.). inid ancestor. These similarities are not meant to (3) Another morphological feature that imply, however, that there are no differ- is not found within the australopithecines ences between G. bilaspurensis and Aus- is the posteriorly diverging tooth row of tralopithecus. There appear to me to be G. bilaspurensis. This condition demon- four main differences, three of which are strates the undeniable links of the Bilas- functionally interrelated. (1) The size of pur mandible with late Miocenelearly the canine has been noted above. Simons Pliocene dryopithecines (Pilbeam, ’70), has demonstrated that the angle of oc- but again does not eliminate it from an- clusal wear on the lower canines of G. cestral status. Reduction in molar length bilaspurensis is lower than on the canines and increase in molar breadth accompany- of G. blachi, but higher than on the ca- ing canine reduction would probably pro- nines of Australopithecus (Simons and duce a more parabolic dental arcade. GIGANTOPITHECUS AND AUSTRALOPITHECUS 419 (4) Lower molars of G. bilaspurensis pendence upon tools. But what would lack the complex crenulations of the oc- happen in the Early or Middle Pliocene if clusal surfaces that are so characteristic a hypothetical African G. bilaspurensis of Australopithecus. These teeth tend to began to use tools? First, the large body be long compared to breadth (see table 3), size which could be construed as a defen- but are not without parallels in the aus- sive mechanism could become smaller. tralopithecines. Simons and Chopra (‘69a) Canines would reduce in size in respond- point out that the molars are unlike apes ing to tool manipulation or possibly in in that they have low broad cusps, and response to Jolly’s “phase I” adaptation that they seem to more “closely parallel (‘70). Whatever the case, with smaller hominids and perhaps Pongo” (’69a:13). canines G. bilaspurensis or something In summary, the major differences be- similar to it would look very much like an tween Australopithecus and G. bilaspuren- ER-729 or ER-818. The mandible would sis are the absolute size, wear, and mor- lose anterior robusticity due to the smaller phology of the canine, the polycuspid canines, and length reduction and breadth lower third premolar, the posteriorly di- expansion of cheek teeth related to selec- verging dental arcade, and the lack of tion pressures of heavy mastication would heavily crenulated but mesio-distally long result to counteract interstitial wear. molars. All but the last are related pri- From table 3 it can be seen that G. marily to the anterior and sug- bilaspurensis has a larger LIB index for gest a complex not unlike Ramapithecus, MI, MB, or M3 than any G. blacki or the and, in light of the other similarities, not mean of either robust or gracile australo- particularly crucial to the decision to pithecines. Reduction of mesio-distal length accept or reject it as a possible hominid and increase in buccal-lingual breadth ancestor (see Andrew’ reasons for retain- is reasonable when considering the amount ing Ramapithecus, ’71). A minimum of interstitial wear of both Australopithe- amount of selection to reduce canine size cus and Gigantopithecus (Wolpoff, ’71b). could very plausibly produce an East By increasing buccal-lingual breadth, the Rudolf-like australopithecine from some- amount of occlusal area lost by interstitial thing like G. bilaspurensis. wear would be considerably less, and, hence, selectively important. Coupled DISCUSSION with this, reduction in Ps-M3 length would Recently, Eckhardt (‘71, ’72) has pro- be the mechanical advantage of a shorter posed that the genus Gigantopithecus mandible with more forward placement could be ancestral to hominids. His main of m. temporalis and m. masseter (An- argument concerning the amount of time drews, ’71). With shorter P3-M3 length, required for morphological change may be canine reduction and greater breadth of valid, but the inclusion of G. blachi as an all the cheek teeth, a parabolic arcade ancestor is certainly debatable. From all would be produced. Both G. blachi and indications, G. blacki is no earlier than Australopithecus demonstrate this change. the late Pliocene and probably as recent In each the form of the tooth row is un- as Middle Pleistocene. In all probability it questionably parabolic (see Eckhardt, overlaps with Australopithecus from Afri- ’72: 103). ca, and very likely overlaps with Homo One of the most crucial questions con- erectus (Pei, ’60; Kahlke, ’61; von Koenigs- cerning the relationship of G. bilaspuren- wald, ’52; Woo, ’62). Because of this time sis to Australopithecus is the sex of the overlap, G. blachi cannot be considered Bilaspur mandible. Previous studies have ancestral to australopithecines. sexed the mandible as female (Simons G. bilaspurensis, however, is consid- and Chopra, ’69a: p. 6; and Simons and erably earlier (at least Late Miocene) Ettel, ’70: p. 79). I submit that the sex of (Simons and Chopra, ’69a,b; Simons and a single specimen which is the sole rep- Pilbeam, ’72) and as has been demon- resentation of a new species cannot be ac- strated, is metrically and morphologically curately assessed beyond the 50% level, similar to australopithecines. It is de- i.e., a guess. The Bilaspur mandible is no scribed by most authors as a pongid oc- less likely a male, and, if it is, one would cupying a savannah niche without de- expect to find smaller females which would 420 DAVID W. FRAYER

TABLE 3 Lengthlbreadth indices for lower molars of G. blacki, G. bilaspurensis, and Australopithecus. All indices are multiplied by 100. Bracketed numberfollowing mean refers to sample size. All dental measurements for Australopithecus are listed in Appendix

Mi Mz M3 G. blacki I 106.0 106.9 - G.blacki I1 116.9 112.7 - G. blacki 111 102.2 101.0 113.8 G.bilaspurensis IV 119.7 119.4 121.0 Australopithecus (robust) Mean 102.1 (28) 106.1 (29) 114.2 (17) s.d. = 6.8 s.d. = 5.1 s.d. = 5.9 Range (86.1-113.7) (90.0-116.3) (101.8-123.0) Australopithecus (gracile) Mean 106.2 (23) 106.8 (22) 108.9 (11) s.d. = 9.3 s.d. = 7.6 s.d. = 10.9 Range (80.8-119.3) (87.S119.2) (94.8-1 34.7) even more closely resemble australopithe- though, with effective hunting technology cines. could have caused extermination, as could In the middle and late Miocene there have competition with the were, at least, two possible hominid ances- (Simons, '72). The late existence of these tors, Ramapithecus and G. bilaspurensis. large proves their effective adap- Until now, no one has recognized the pos- tation through size. sibility of the species represented by the As a hypothesis, this suggestion will Bilaspur mandible as being ancestral to stand or fall as new specimens are dis- early hominids. I feel that G. bilaspurensis covered. It is important to realize that the is a better candidate for an australopith- issue of man's Pliocene ancestry is far ecine forbear in that body size is an im- from closed, and that the present evidence portant adaptive mechanism in creatures does not yet allow a clear decision between that are exploiting a terrestrial-niche Ramapithecus and G. bilaspurensis. It is without dependence on tools. Livingstone important that viable alternative hypoth- ('62) as well as Weidenreich ('45) have eses be considered as additional evidence previously suggested the importance of is discovered. size and reduction of size in human evo- ACKNOWLEDGMENTS lution. G. bilaspurensis closely follows an aus- The original version of this paper was tralopithecine morphological pattern in discussed with Dr. Elwyn L. Simons (Yale both metric and non-metric characteris- University), and, although we disagree, tics. Its only major differences are not I truly appreciate the time he spent with crucial in eliminating it as a hominid me. Drs. Milford H. Wolpoff and C. Loring ancestor since Ramapithecus evinces the Brace (University of Michigan) have both same morphological differences, and in critically commented on this version, as many other respects, is more apelike. has Dr. Alan Walker (University of Nairo- Gigantopithecus blacki, on the other bi). Tooth and mandibular measurements hand, can best be explained as an Asiatic compiled by Wolpoff and Mr. Paul Mahler hominoid (an aberrant hominid) which are gratefully acknowledged. The inter- did not use tools, but rather continued to pretations and conclusions of this paper increase in body size as it adapted to the are those of the author. forest conditions. From the number of LITERATURE CITED teeth found in China, it would seem that Andrews, P. 1971 Ramapithecus wickeri man- this specific adaptation (size 'us tools) was dible from Fort Ternan, Kenya, , 231: quite adequate. Survival into the mid- 192-194. Pleistocene attests to this. Australopithe- Bishop, W.W., and G. R. Chapman 1970 Early cus with smaller size and a non-hunting Pliocene sediments and fossils from the northern Kenya Rift valley. Nature, 226: 914-918. industrial technology could not drive G. Dart, R. 1926 Australopithecus africanus: The blachi into extinction. , man- of S. Africa. Nature, 1 15: 195-199. GIGANTOPITHECUS AND A USTRALOPZTHECUS 421

Eckhardt, R. B. 1971 Hominoid Dental Varia- Pei, W. C. 1960 The living environment of tion and Human Origins. Doctoral dissertation, Chinese primitive men. Vert. Pala., 4: 4044. University of Michigan. Pilbeam, D. 1970 Gigantopithecus and the 1972 Population genetics and human origin of . Nature, 225: 516-519. evolution. Sci. Am., 226: 94-103. Simons, E. L. 1972 Evolution. Mac- Howell, F. C. 1969 Remains of Hominidae from millan Co., New York. Pliocene/Pleistocene formations in the lower Simons, E. L., and S. R. K. Chopra 1969a Gi- Omo basin, Ethopia. Nature, 223: 1234-1239. gnntopithecus (Pongidae, Hominoidae) a new Jolly, C. J. 1970 The seed-eaters: a new model species from North India. Postilla, 138: 1-18. of hominid differentiation based on a baboon analogy. Man, 5: 1-26. 196913 A preliminary announcement Kahlke, H. D. 1961 On the complex of - of a new Gigantopithecus species of India. In: Ailuropoda fauna of Southern China and the Recent Advances in Primatology. Vol. 2. H. 0. chronological position of Gigantopitheczis blacki Hofer, ed. Kargor, Basel, pp. 135142. von Koenigswald. Vert. Pala., 2: 104-108. Simons, E. L., and P. C. Ettel 1970 Giganto- von Koenigswald, G. H. R. 1952 Gigantopithe- pithecus. Sci. Am., 222: 77-85. cus blacki von Koenigswald, a giant fossil hom- Simons, E. L., and D. Pilbeam 1965 Prelim- inoid from the Pleistocene of Southern China. inary revision of the Dryopithecinae (Pongidae, Anthro. Papers of Am. Mus. of Natl. Hist., 43: Anthropoidea). Folia Prim., 3: 81-152. 291-325. 1972 Hominoid paleoprimatology. In: 1958 Gigantopithecus and Australo- The Functional and Evolutionary Biology of pitkecus. The Leach, 28. Primates. Russell Tuttle, ed. Aldine, Chicago, Leakey, R. E. F. 1971 Further evidence of Low- pp. 36-62. er Pleistocene hominids from East Rudolf, North Weidenreich, F. 1945 Giant early man from Kenya. Nature, 231 : 241-245. and South China. Anthro. Papers of Am. 1972 Further evidence of Lower Pleis- Mus. of NatI. Hist., 40: 1-134. tocene hominids from East Rudolf, North Kenya, 1946 Apes, Giants, and Man. Univer- 1971. Nature, 237: 26G269. sity of Chicago Press, Chicago. Leakey, R. E. F., and J. M. Mungai and A. C. 1949 Facts and speculations concern- Walker 1971 New australopithecines from ing the origin of Homo sapiens. Anthro. Papers East Rudolf, Kenya. Am. J. Phys. Anthrop., 35: of Franz Weidenreich, Viking Fund, New York. 175-186. Wolpoff, M. H. 1971a Metric Trends in Hom- 1972 New australopithecines from inid Dental Evolution. Case Western Reserve East Rudolf, Kenya (11.) Am. J. Phys. Anthrop., University Press, Cleveland. 36: 235-252. 1971b InterstitiaI wear. Am. J. Phys. Livingstone, F. B. 1962 Reconstructing man’s Anthrop., 34: 205-228. Pliocene pongid ancestor. Amer. Anthro., 64: Woo, T. K. 1962 The mandibles and dentition 301-305. of Gigantopithecus. Paleo. Sinica, 12. 422 DAVID W. FRAYER

APPENDIX 1 Mandibular dimensions for Australopithecus (gracile and robust) and Gigantopithecus. Except where noted, all measurements are from the original specimen. Abbreviations signify the site or area where the fossil was discovered. (SK, Swartkrans; STS, Sterkfontein; MLD, Maka- pansgat; TM, Kromdraai; Sterkfontein; ER, East Rudolf; OH, Olduvai). (Inmm)

External Maximum Corpus Corpus Specimen bicanine symphyseal height breadth Length breadth length at MI at Mz p4-M~ Australopithecus (robust) TM-1517 33.8 28.3 39.4 SK-12 31.0 50.0 41.0 34.0 40.0 SK-23 32.8 48.5 35.1 27.8 39.4 SK-34 41.0 22.0 43.1 SK-74 36.0 37.8 SK-858 37.0 40.7 Omo-7-125f 26.0 1 50.7 1 47.0 32.7 44.7 ER-403 46.6 31.8 42.8 ER-404 48.3 35.8 ER-725 40.0 31.4 42.5 ER-726 44.5 29.5 51.0 ER-728 37.5 25.8 ER-729 36.1 51.5 45.7 29.5 47.5 ER-810 38.0 27.3 44.0 ER-818 51.0 36.0 50.8 Natron 35.0 50.6 36.7 31.5 46.0 Australopithecus (gracile) MLD-18 35.2 32.4 25.4 35.4 MLD-40 35.4 29.5 38.0 STS-7 45.0 36.0 37.5 STS-36 36.0 23.4 STS-52 36.7 39.5 ER-730 31.7 32.7 33.2 18.8 32.1 ER-992 38.5 34.0 23.2 34.6 Gigantopithecus G. blacki I 45.3 2 76.0 60.0 30.0 50.5 G. blacki I1 - 77.0 53.0 31.0 3 57.7 G. blacki 111 56.7 2 108.5 81.0 34.0 57.1 G. bilaspurensis 36.0 2 62.4 2 52.1 2 34.2 2 49.2

1 From Howell ('69) or from cast belonging to Howell. 2 Measured from casts produced by Yale University. 3 Behind Pq. GIGANTOPITHECUS AND A USTRALOPITHEC US 423

APPENDIX 2

Dental measurements of mandibular teeth 2 of gracile australopithecines. All measurements are from the original specimens unless otherwise indicated. (In mm)

Incisor 2 Canine Premolar 3 Premolar 4 Molar 1 Molar 2 Molar 3 Specimen 1 b1 b1 b 1 b 1 b 1 b 1 b

Omo-75 1 11.3 12.3 11.4 12.7 15.6 14.1 17.5 15.4 15.1 14.1 Omo-Kalam 7 1 15.1 13.4 Omo-75 S.15 1 14.0 13.0 Omo-WS-508 1 13.3 12.2 Omo-WS-752 1 i4.1 13.0 Omo-L2-89 1 13.7 11.5 Omo-L45-2 1 12.9 12.0 Omo-L51-1 1 14.0 12.6 Omo-L26-1 15.5 13.0

Omo-L28-30131 1 15.0 13.0 ~.16.7 12.4~~ ~ OH-7 7.2 7.4 8.9 9.8 9.5 10.2 10.3 10.6 14.1 12.5 15.8 13.8 OH-16 7.6 7.6 9.9 10.1 10.3 11.5 10.1 11.0 14.3 12.8 15.3 14.7 15.8 14.4 OH-30 6.8 7.8 7.7 16.5 14.5 ER-730 11.3 12.0 11.6 11.8 13.0 11.6 ER-992 7.1 7.0 9.0 9.2 9.3 11.5 8.6 11.4 12.3 10.9 12.8 12.3 13.1 12.5 STS-4 14.4 13.3 STS-6 14.3 14.0 STS-7 5.5 8.5 9.2 11.0 10.9 13.2 10.7 12.5 14.3 14.0 15.0 14.6 15.3 14.7 STS-9 14.9 13.0 STS-18 15.3 14.5 STS-24 7.6 6.5 13.6 11.4 STS-36 9.5 10.5 9.5 13.3 9.0 13.3 11.8 14.6 14.8 16.9 16.3 17.2 STS-52b 7.0 8.1 7.9 10.2 9.1 11.9 10.0 11.6 13.5 13.1 14.5 13.4 13.8 12.7 STS-55b 15.2 13.3 TM-1515 9.0 13.0 10.4 13.0 16.9 16.4 TM-1518 14.3 13.2

Taung 16.9~~~ 16.4~~~ MLD-2 10.0 12.6 11.1 14.7 14.1 16.8 15.1 MLD-18 5.8 8.0 8.9 8.9 8.3 11.4 8.4 12.0 12.4 13.1 14.6 14.8 14.0 13.9 MLD-22 13.5 14.3 15.8 15.2 MLD-24 15.0 14.0 MLD-29 9.0 12.0 12.5 13.4 MLD-40 7.7 9.1 8.9 11.4 9.5 11.4 12.8 12.3 15.0 13.8 16.5 14.1

1 From Howell ('69). 2 Only isolated MI and MZ are included. 424 DAVID W. FRAYER

APPENDIX 3

D(wta1measurements of mandibular teeth 2 of robust nitstmlopithecines. All measurements are from original specimens unless otherwise indicated. (In mm)

Incisor 2 Canine Premolar 3 Premolar 4 Molar 1 Molar 2 Molar 3 Specimen lblblblblblblb

Omo-L7-125 1 7.8 9.6 10.4 17.5 11.7 18.9 16.8 18.7 16.2 18.0 18.2 14.8 Omo 74 1 8.8 9.7 13.0 13.8 Natron 6.0 6.3 7.5 8.0 9.3 13.4 14.0 15.0 15.4 15.3 17.0 16.1 18.0 16.0 TM-1517 9.4 8.8 10.3 12.7 10.8 12.9 13.8 13.2 15.4 14.4 16.4 14.0

TM-1536 12.5~ ~~~11.8 TM-1600 9.9 12.1 15.0 14.8 15.9 14.8 SK-1 17.0 15.3 SK-5 14.9 14.2 SK-6 13.0 10.3 10.7 12.9 15.9 15.4 16.9 16.3 18.5 15.4 SK-12 10.0 12.0 10.5 12.0 15.0 14.4 15.8 15.8 16.0 15.2 SK-15 12.0 12.0 13.0 12.7 14.3 12.7 SK-20 14.7 14.2 SK-23 7.0 6.6 8.2 8.0 9.1 11.3 10.0 14.2 14.5 14.6 14.9 14.8 16.0 13.2 SK-25 11.4 13.2 14.7 14.0 16.3 14.7 SK-34 6.6 7.0 8.9 8.8 9.7 12.8 13.0 14.0 13.5 14.5 16.6 16.3 17.0 16.7 SK-37 16.5 14.7 SK-45 11.3 12.6 12.3 SK-55 10.0 10.3 14.6 13.7 16.1 14.3 15.5 13.8 SK-61 15.0 14.0 SK-63 13.7 13.5 SK-74 8.1 7.6 9.2 8.8 10.1 10.1 12.1 13.2 13.5 14.5 14.3 SK-81 9.1 10.6 10.6 13.3 14.3 15.8 16.0 16.0 17.0 15.0 SK-104 14.9 13.9 SK-828 15.4 14.2 SK-838b 14.1 13.0 SK-843 14.4 13.4 15.9 14.0 17.3 15.0 SK-846a 14.6 13.7 SK-858 6.7 7.5 8.5 8.0 10.0 13.7 11.0 13.9 15.0 15.8 15.0 SK-876 9.0 10.8 10.0 11.7 10.5 12.5 13.0 15.1 17.0 15.1 18.5 15.7 SK-1586 13.7 15.0 14.0 16.5 15.0 SK-1587 10.4 11.0 13.3 13.0 15.0 12.9 SK-1588 10.5 11.6 14.1 12.4 SK-1648 12.5 15.7 14.6 SK-3974 14.8 13.5 SK-3976 17.4 16.0 ER-729 8.5 10.1 11.4 13.2 14.2 14.5 15.6 15.7 19.0 18.0 20.5 18.7 ER-801 13.8 14.5 18.1 16.5 18.8 15.5 ER-810 17.4 15.6 ER-818 13.4 13.9 15.1 16.4 15.2 18.5 18.0 21.5 18.4 ER-1171/2 18.8 16.9

1 From Howell ('69). 2 Only isolated MI and MZ are included. GIGANTOPITHECUS AND A USTRALOPlTHECUS 425

APPENDIX 4 Mandibular dimensions of P. gorilla (male and female). Specimen numbers refer to individual catalogue numbers in the Cleveland Museum of Natural History. (In mm)

External Maximum Corpus Corpus Specimen bicanine symphyseal height breadth Length breadth length at M2 at Mz p4-M~ P. gorilla (male) B-647 51.5 59.7 31.7 17.9 43.2 B-1076 60.5 63.2 39.7 20.7 48.7 B-1402 55.2 62.0 35.8 20.2 45.1 B-1404 56.7 73.3 40.6 18.6 45.6 B-1405 62.4 68.0 38.9 20.0 40.3 B-1731 54.7 65.2 39.6 19.6 40.3 B-1736 61.0 75.2 40.8 19.8 40.9 B-1746 62.3 72.2 38.9 22.1 44.3 B-1784 54.6 72.0 40.7 18.6 46.3 B-2000 57.9 76.2 43.2 22.8 43.9 B-2029 56.8 69.1 37.1 19.9 43.2 B-2766 61.6 69.1 38.2 21.0 46.5 B-2767 56.5 74.6 43.2 20.8 46.2 B-2826 62.2 75.4 40.4 23.6 46.4 B-3410 52.3 68.4 33.6 21.1 46.1 B-3415 61.3 62.0 39.6 19.9 45.8 B-3420 57.0 72.2 38.9 27.7 50.0 B-3431 54.5 62.2 36.5 18.4 40.6 B-3547 51.9 69.9 32.7 21.4 40.3 B-3557 59.1 65.7 39.3 25.1 44.9 P. gorilla (female) B-1398 43.1 56.6 35.8 21.5 41.1 B-1399 50.5 55.9 33.9 22.0 44.2 B-1400 46.0 58.9 38.1 23.0 43.3 B-1419 45.6 50.7 31.8 21.5 42.2 B-1690 47.3 57.4 32.8 23.9 43.5 B-1710 46.0 49.5 32.6 21.4 35.9 B-1725 50.3 54.8 34.1 26.8 42.0 B- 1756 49.0 53.4 31.9 24.4 39.6 B-1851 47.6 53.4 35.4 26.8 41.2 B-1996 47.8 53.2 30.2 19.5 43.5 B-2782 43.7 51.7 32.4 20.3 44.8 B-2785 47.6 54.0 34.0 19.3 44.6 B-2799 42.6 50.1 32.1 18.8 44.4 B-2818 44.0 54.6 31.4 21.7 41.4 B-2820 46.8 49.0 32.0 18.7 41.0 B-3393 45.9 51.2 29.4 16.8 40.7 B-3405 49.0 53.8 29.7 24.8 38.9 B-3424 43.6 47.6 33.5 18.7 40.0 B-3426 44.0 50.3 27.0 17.3 40.4 B-3562 48.7 57.8 32.8 20.2 42.9 APPENDIX 5 Dental dimensions ofmandibular teeth of male and female P. gorilla. All specimens are .from the Cleveland Museum of Natural History. (Inmm) Incisor 1 Incisor 2 Canine Premolar 3 Premolar 4 Molar 1 Molar 2 Molar 3 Specimen 1b 1b 1b 1b 1b 1b 1b 1b Male B-647 9.2 8.9 10.2 16.3 13.3 15.4 10.8 B-1076 10.0 8.8 9.9 18.8 15.5 18.1 11.7 10.9 12.8 15.3 13.0 B-1402 9.8 9.9 10.9 16.3 13.2 18.1 11.4 12.4 13.8 17.2 14.2 8.5 11.7 16.2 14.6 17.0 15.2 16.2 14.5 B-1404 9.3 19.0 13.8 17.6 12.2 12.1 13.5 16.9 14.8 B-1405 8.8 7.2 8.5 18.1 12.6 15.1 10.9 13.0 14.0 15.5 12.8 19.1 16.7 7.5 9.6 13.6 12.0 16.8 16.5 17.0 15.8 B-1731 7.4 8.4 17.1 13.6 16.0 12.5 11.0 12.5 8.2 9.9 10.5 12.8 14.2 13.0 17.1 15.2 18.5 15.3 8.7 8.5 9.8 10.8 18.1 14.1 17.0 11.7 15.0 16.8 14.7 9.0 18.9 15.0 10.5 12.6 14.8 13.5 15.718.1 14.4 B-1736B-1746 8.6 11.5 9.9 18.5 12.6 11.5 13.7 14.7 13.2 17.8 14.8 16.9 14.6 6.7 9.1 9.5 19.2 15.5 17.015.6 14.913.3 15.8 14.3 B-1784 - - 17.9 11.7 12.4 13.9 16.1 13.5 15.6 14.1 6.2 9.5 9.3 17.5 14.1 17.8 12.2 11.9 13.8 15.0 13.3 17.1 14.9 17.2 15.1 B-2000 5.1 16.3 13.2 16.1 14.2 8.4 10.0 12.0 16.0 10.3 11.0 11.8 15.1 13.4 18.4 15.0 7.1 9.6 __ 20.8 16.0 18.0 16.7 17.9 15.3 B-2766B-2029 19.3 14.3 18.1 11.7 12.2 14.1 15.9 13.9 17.0 14.2 8.3 9.3 9.0 11.1 12.0 14.5 16.2 14.4 18.2 15.6 ?i B-2767 8.1 8.5 12.0 20.6 15.1 17.8 12.8 16.518.0 15.615.1 @ 9.3 19.5 14.6 11.9 15.0 16.3 14.1 18.0 16.5 B-2826 -8.3 -9.9 9.0 11.0 18.8 13.1 18.0 15.4 18.918.1 14.616.4 18.9 12.9 12.1 14.1 16.0 14.0 8.3 9.8 11.5 18.2 14.7 15.6 13.8 20.0 18.7 19.718.4 17.815.5 5 B-3415B-3410 8.7 9.0 9.5 20.0 17.1 17.7 12.4 12.2 13.0 16.0 13.9 16.0 13.6 7.0 9.8 8.0 9.6 19.0 14.8 21.6 12.6 12.710.0 15.312.4 17.3 15.0 16.0 14.2 z 7.3 10.3 12.0 14.3 12.9 15.7 14.1 B-3420B-3431 7.0 9.0 8.0 10.4 16.5 12.9 16.8 11.6 11.1 14.4 14.6 13.1 17.9 16.1 9.c 8.8 17.0 11.5 18.0 16.1 7.7 19.8 16.0 15.8 14.5 M B-3557 18.5Female 13.0 5 B-3547 7.8 8.6 8.1 9.3 13.5 9.8 10.3 11.0 16.5 14.2 5.17.7 8.9 8.1 9.1 12.0 9.2 14.2 10.2 11.8 11.8 16.2 14.5 15.9 13.2 7.57.2 8.1 9.3 9.4 13.8 9.6 11.2 11.4 14.6 12.6 16.6 14.4 11.8 10.0 15.3 12.4 17.1 13.8 7.2 8.1 8.6 9.2 12.7 10.2 15.2 10.4 10.8 12.9 16.5 14.3 16.4 14.1 B-1398 7.0 8.9 8.2 11.0 16.0 11.2 11.1 13.0 15.6 13.1 17.4 15.6 B-1399 - 7.5 8.8 13.7 9.8 9.2 11.2 15.1 13.2 16.3 14.4 6.7 7.7 14.5 9.4 14.7 12.8 7.8 10.8 13.9 10.5 11.2 14.2 15.5 13.3 16.9 15.2 B-1400 6.8 9.1 12.8 10.0 17.2 11.8 13.8 13.2 15.8 14.9 B-1419 7.2 7.3 7.8 13.7 10.7 14.7 10.1 10.2 11.9 12.9 11.5 15.1 13.7 B-1690 5.7 6.9 8.9 10.8 12.8 16.2 15.2 -5.6 8.7 11.9 10.0 15.0 11.0 14.6 13.8 15.2 13.5 15.8 13.7 B-1710 9.9 10.0 11.0 13.1 16.0 14.7 8.8 8.6 10.2 10.3 14.7 10.7 15.7 11.2 14.2 12.0 16.2 15.1 7.3 8.3 11.1 13.7 14.2 13.3 16.0 15.0 B-1725B-1756 6.6 8.5 10.5 13.6 10.1 15.1 12.0 17.0 15.6 B-1851 8.5 8.8 10.1 9.4 14.6 10.6 15.6 11.2 11.8 14.2 15.5 13.6 17.9 15.8 17.6 15.5 8.0 8.8 9.6 9.0 14.9 9.9 11.7 11.3 15.8 14.4 15.5 13.2 B-1996 14.0 11.0 15.5 13.5 17.3 16.2 15.3 13.4 B-2782 5.9 7.9 7.1 8.3 13.5 11.1 13.8 10.3 10.5 12.7 17.9 14.3 7.8 8.3 9.1 10.3 11.8 14.8 12.9 15.9 14.0 B-2785 7.5 12.5 10.3 13.5 9.9 14.8 13.2 16.1 13.8 14.6 13.0 B-2799 5.8 8.0 7.2 9.1 14.4 9.7 10.3 12.1 16.0 13.6 8.3 12.7 10.3 9.6 12.0 14.7 12.7 15.7 15.5 B-2818 7.0 -8.4 - 12.3 -9.4 14.0 10.5 14.7 12.8 15.7 14.4 8.9 10.1 13.0 10.7 9.8 13.0 16.0 13.3 B-2820 7.3 12.8 8.8 14.9 10.3 13.5 12.2 15.8 13.3 8.1 9.0 9.8 9.5 13.2 10.6 14.2 9.4 10.4 12.7 16.0 14.3 B-3393 13.0 15.0 11.0 10.9 12.8 14.6 13.2 15.6 13.3 15.3 13.9 8.5 14.5 12.6 15.5 13.9 B-3405 14.8 13.1 17.2 14.5 B-3424 B-3426 B-3562