Cranial remains of Middle Pleistocene European hominids

Milford H. Wolpoff Department of Anthropology, University of Michigan, Ann Arbor, Michigan 48104, U.S.A. Received 30 January 1979 and accepted 14 May 1979 Keywords: evolution, sexual dimorphism, Steinheim, Swanscombe, Petralona, Bilzingsleben, V6rtessz6116s, Arago, La Chaise, Biache.

Cranial R e m a i n s o f Middle P l e i s t o c e n e European Hominids Cranial remains of the Middle Pleistocene European hominids are reviewed in a historical context. Swanscombe and Steinheim, the earliest specimens to be discovered, are compared with each other. Following this, the more recently discovered specimens from V6rtessz6116s, Petralona and Bilzingsleben are compared. Finally, Riss remains from La Chaise, Biache and Arago are discussed in the context provided by the above specimens. It is suggested that while phyletic evolution may play a role in the observed differences, a marked degree of sexual dimorphism contributes to the variation expressed by these specimens.

1. I n t r o d u c t i o n T h e Middle Pleistocene hominids of Europe represented by cranial remains are usually given fairly cursory treatment in discussions of h u m a n evolution. T h e best known specimens, Steinheim and Swanscombe, were described decades ago. While alleged differences between these were once a focal point for the presapiens hypothesis, most workers have come to recognize their similarities as well as the futility of debating what the supraorbitals of Swanscombe m a y or m a y not have been like. Discussions of European hominid evolution usually move quickly on to the origin and fate of the Neandertals. T h e more recently discovered specimens have not been fully described (excepting V6rtessz6116s). T h e y have never been systematically compared with each other, or with Steinheim and Swanscombe. Indeed, a systematic comparison of the latter two has never been attempted. T h e total sample size available has become larger; the latter portions of Middle Pleistocene hominid evolution are probably better represented in Europe than anywhere else. While the sample bears on the problem of U p p e r Pleistocene hominid origins in Europe, other questions involving the extent of sexual dimorphism a n d its evolution, the nature of the Homo erectus-Homo sapiens transition, the identification of functionally important morphological complexes, and the clarification of European evolutionary trends await treatment of the sample as a whole. This p a p e r will review the Middle Pleistocene European cranial sample, based on a recent examination by the author of all the specimens in this sample but constrained in some eases by the limitations of what has been published thus far. T h e specimens are discussed in a historical context, beginning with the earliest discoveries at Steinheim a n d Swanscombe. Steinheim is compared with the less complete British specimen, and these in turn are systematically c o m p a r e d with the late Homo erectus sample from Choukoutien, a n d with the succeeding European Neandertals. T h e later discoveries from V6rtessz6116s, Bilzingsleben and Petralona are then treated in the same manner, and finally a brief comparison is m a d e with the Riss specimens from L a Chaise, Biache a n d A r a g o . T h e effects of phyletic evolution a n d sexual dimorphism on the metric and morphological variability observed, and the apparent European evolutionary trends leading to the U p p e r Pleistocene hominids, are discussed in the concluding portions of the paper. 2. S t e l n h e i m a n d S w a n s c o m b e These were the first of the early European crania to be discovered. Steinheim was found Journal of Human Evolution (1980) 9, 339-358 0047-2484/80/050339 + 23 $02.00/0 (~ 1980 Academic Press Inc. (London) Limited

340

~. H, WOLPOFF

in a gravel pit in the vicinity of Steinheim an der Murr, near Stuttgart. The announcement was made by Berckhemer (1933) and a fairly accurate, although far from complete description, was published by Weinert several years later (1936). Swanscombe was also recovered in a gravel pit, in this case from Kent in Great Britain. A parietal and occiput found in 1935 and 1936 were described exhaustively in works by Marston (1937), Le Gros Clark (1938) and M o r a n t (1938). The unlikely but fortunate discovery of the other parietal in 1955 led to a new analysis of the vault by a number of workers which was compiled in a volume edited by Ovey (1964). For over 30 years, these provided the only cranial information regarding the origin and early evolution of Homo sapiens in Europe. When F. C. Howell reviewed the Middle Pleistocene European hominids in 1960, Steinheim and Swanscombe were still the only specimens known with cranial remains that he felt could be dated to the Mindel-Riss interglacial. At the time, these specimen were pivotal in the presapiens hypothesis as presented by Vallois (1954, 1958) and Heberer (1950) since they were thought to represent different European lineages. Howell [as well as Morant (1938), Breitinger (1955), Le Gros Clark (1955) and others] had recognized a fundamental similarity between these two crania. However, he used them to buttress an argument supporting a lineage division between these Europeans and (what he called) contemporary peoples of eastern Asia. Such differences in phylogenetic reconstruction are hardly surprising in view of the various interpretations that were placed on the specimens themselves. These ranged from claims of no fundamental difference from modern people (Weidenreich, 1940) to more similar to modern Homo sapiens than to Neandertals (Le Gros Clark, 1955), and finally to a claim of intimate relationship with other archaic Homo sapiens specimens (Sergi, 1953 ; Roginskii, 1948). As the result of my recent examination of these specimens, I believe that two broad contentions concerning their relations to each other and to other Homo sapiens samples can be supported. First, the specimens are indeed similar where they can be directly compared. Second, insofar as they differ from the later Neandertals, they either are more like Homo erectus or intermediate between Homo erectus and Neandertals. In no aspect of the total morphological pattern that can be observed are either of these crania more like anatomically modern Homo sapiens than are the Neandertals that followed them in time. Regarding the first point, it should be recognized that direct comparisons of the crania are limited to the parietals and occipital, and constrained by the amount of warping and distortion that Steinheim has suffered (Plate 1). I believe Howell (1960, p. 216) was incorrect in stating that the right side "is perfectly preserved" and that the distortion was "wrongly emphasized" by Weinert (1936). While it is true that most of the morphological features are retained on this side, and that the bone surface is largely continuous, there has been marked medial crushing of the occiput, possibly also involving the mastoid region of the temporal. A line taken from the palate mid.line to basion passes well laterally of the opisthion postion (Plate 2). Any measurements of Steinheim must be regarded with caution. M y attempts to obtain dimensions for this cranium utilized symmetric reconstruction from the undistorted portions of the right side, or unambiguous compensations for distortion. No measurements involving obvious distortion that could not be compensated for were attempted. A comparison of dimensions (Table 1) reveals that Steinheim is almost invariably the

Plate 1. Comparison of Steinheim (left) and Petralona in lateral, facial and 3/4 views.

340

Plate 2. Comparison of Steinheim (left) and Petralona in basal and superior views.

Plate 3. Some views of the Bilzingsleben frontal (above) and occipital (below) fragments.

MIDDLE PLEISTOCENE EUROPEAN

Table 1

HOMINIDS

341

Vault measurements that can be compared between Swanscombe and Steinheim, with equivalent data for Petralona and Sal@. A l l m e a s u r e m e n t s were taken by the author on the original specimens (or in the case of Sal~ an accurate cast) and are in millimeters. In this and all other tables, when measurements could be taken of both sides of the specimen~ the average is given

Steinheim* Swanscombe Petralona

Sal6++

Biasterionic breadth 102.5 t 121.8 119.0 116.0 Lambda-inion chord 60.4 58.0 63.3 43.5 Foramen magnum breadth 30-0 30.1 33.9 31.5 Biparietal breadth 126.8 t 145.0 150.0 130.0 Bregma-lambda 91.7 107"5 109 "2 96-0 Sphenion-asterion 83-7 84.0 104.8 87"7 Sphenion-bregma 87-0 89.5 88.0 81 "3 Lambda-asterion 86.7 91.6 87-3 85.1 Basion-bregma 104.5 125.0 127"5 110.0 Basion-lambda 102"0 115.9 115"0 96"0 Basion-inion 80.7 89.5 100.0 83.5 Bregma-inion 136.0 142.0 157.3 128.7 Height (basion-bregma) X breadth (biparietal) x length (bregma-inion) 1.80x 106 2"57× 108 3.01 x 10e 1.84× 10~ * In almost every case these dimensions are less than those published by Howell (1960). I attempted to duplicate the dimensions he recorded and cannot account for the differences. I" Of all the dimensions reported, these (based on symmetric reconstruction from the right side) are the most likely ones to be affected by distortion. ~. Measured on an accurate cast (based on comparisons with published dimensions). s m a l l e r o f the two c r a n i a . T h e difference seems m o s t d r a m a t i c in b i p a r i e t a l a n d occip i t a l b r e a d t h s , a l t h o u g h these are the least c e r t a i n o f the dimensions p r e s e n t e d for Steinheim. T h e p a r i e t a l c o m p a r i s o n s i n d i c a t e t h a t S w a n s c o m b e is s a g i t t a l l y l o n g e r (the v a u l t is m o r e c u r v e d ) a n d slightly m o r e e l o n g a t e d p o s t e r i o r a l l y w h i c h p r o b a b l y c o r r e s p o n d s to the b r o a d e r o c c i p u t . A v o l u m e t r i c e s t i m a t e m a d e b y m u l t i p l y i n g c o m p a r a b l e m e a sures o f v a u l t (or p a r t i a l vault) length, b r e a d t h a n d h e i g h t suggests t h a t S w a n s c o m b e is a p p r o x i m a t e l y 4 0 % larger. T h i s does n o t c o n f o r m to the difference b e t w e e n p r e v i o u s estimates of c r a n i a l c a p a c i t y . H o w e v e r , t h e r e is no g u a r a n t e e t h a t these estimates a r e c o r r e c t or even a p p r o x i m a t e . S w a n s c o m b e , o f course, lacks a frontal, s p h e n o i d a n d t e m p o r a l s . I n the case o f S t e i n h e i m , a p p l i c a t i o n o f the regression f o r m u l a e using the p r o d u c t o f c r a n i a l length, b r e a d t h a n d h e i g h t p r e s e n t e d b y O l i v i e r & Tissier (1975) p r o v i d e r a t h e r different results d e p e n d i n g on w h e t h e r it is c o n s i d e r e d a n e x a m p l e o f " p a l a e o a n t h r o p i i " (loosely N e a n d e r t a l ) o r a n " a r c h a n t h r o p i i " (Homo erectus). T h e s e regressions r e s p e c t i v e l y give c a p a c i t y estimates o f 1085 cc a n d 762 cc. H o w e v e r , since S t e i n h e i m fails in n e i t h e r o f these categories it is n o t . c l e a r w h i c h s h o u l d a p p l y . F o r instance, a p p l y i n g the " p a l a e o a n t h r o p i i " f o r m u l a to P e t r a l o n a (see b e l o w ) , a n a p p r o x i m a t e c o n t e m p o r a r y of S t e i n h e i m , results in the u n l i k e l y c a p a c i t y e s t i m a t e o f 1713 cc! Steinheim, in fact, has a s m a l l e r l e n g t h / b r e a d t h / h e i g h t p r o d u c t t h a n a n y Homo erectus

342

M.H. WOLPOFF

s p e c i m e n , b u t e v e n if a m i d w a y e s t i m a t e is used (justifiable b y the fact t h a t the regressions differ o n l y in their i n t e r c e p t values), S t e i n h e i m is m a r k e d l y s m a l l e r t h a n u s u a l l y p r e s u m e d . H o w m u c h s m a l l e r it m i g h t be is suggested b y the N o r t h A f r i c a n c r a n i u m f r o m Sal6 ( J a e g e r , 1975). I n m y view, this c r a n i u m presents n u m e r o u s m o r p h o l o g i c a l similarities to b o t h S w a n s c o m b e a n d S t e i n h e i m . T h e d i m e n s i o n s o f Sal6 a r e s i m i l a r to those o f S t e i n h e i m (where c o m p a r a b l e , see T a b l e 1), a n d the c r a n i a l c a p a c i t y r e p o r t e d for Sal6 is 930--960 cc. A p a r t f r o m the size difference, the m o r p h o l o g i c a l features a n d g e n e r a l p r o p o r t i o n s o f the S t e i n h e i m a n d S w a n s c o m b e c r a n i a a r e s i m i l a r w h e r e c o m p a r a b l e . T h e a p p l i c a b l e features w h i c h Vallois (1954) used to distinguish the " p r e s a p i e n s " E u r o p e a n l i n e a g e i n c l u d e g r e a t v a u l t thickness, p a r i e t a l o f the " H o m o sapiens" type, p r i m i t i v e a r r a n g e m e n t o f t h e p a r i e t o - t e m p o r a l a r t i c u l a t i o n , a n d a n u n b u n n e d o c c i p u t w i t h a h i g h a n g l e for the o c c i p i t a l p l a n e . M o s t o f these a r e s i m i l a r if n o t v i r t u a l l y i d e n t i c a l in these c r a n i a . Both c r a n i a l a c k a n y significant d e v e l o p m e n t o f the n u c h a l torus. S t e w a r t (1964) d e s c r i b e d the N e a n d e r t a l - l i k e m o r p h o l o g y o f the o c c i p i t o m a s t o i d torus o f S w a n s c o m b e , Table 2

M e a s u r e m e n t s o f p a r i e t a l a n d o c c i p i t a l t h i c k n e s s f o r Chouk o u t l e n ( f r o m W e l d e n r e l c h , 1943) a n d f o r E u r o p e a n M i d d l e a n d U p p e r P l e i s t o c e n e a r c h a i c H o m o sapiens s p e c i m e n s m e a s u r e d b y t h e a u t h o r (in r a m )

Parietal thickness at "~ ~ Occipital Middle Mastoid thickness Bregma eminence angle at inion Choukoutien (av.) 3 5 10 11 12

8"6

12.1

15"1

15"5

9.6 9.0* 7.5 7.0 9.7

11.0 12-5 16.0 9.0

18.0 15.0 14.0 13.5 15-0

20-4 15.0 15"0 12.0 15.0

Bilzingsleben Steinheim Swanscombe

9.6* 6.0 7.3

6.5 11.5

6.5 10.1

17'0 11.1 9-0

Neandertal (av.) Vindija 204 205 Spy 1 Spy 2 Le Moustier Wildscheuer A Monsempron Kl~ma Mr. Circeo Saccopastore 2 La Quina 13 La Chapelle La Ferrassie 1 Gil~raltar Neandertal La Quina 5

6"3

8"3

7.1

11.4

8.3 7"0 7.0 6.0

6-0 6-5 5"5

8"0 6-8 9-0 9"0 11-3 7"0 7.1 8.3 8.0 10.0 10.5 5"0

* Frontal thickness at bregma is used.

6-0 4.0 7"5

14.0 13.0 13.0

9"7 6.5 8.0 8.35 7"5 8"9 5"0

11.0

10.0 13.0 6"0

343

MIDDLE PLEISTOCENE EUROPEAN HOMINIDS

suggesting that a small mastoid process was present. T h e m o r p h o l o g y of this region is quite similar in Steinheim where a small mastoid process is preserved. Neither c r a n i u m is bunned, a n d Howell (1960) showed that the sagittal c o n t o u r of the occiputs are the same, but for the size difference. T a k e n b y itself, the absence of b u n n i n g m i g h t be interpreted as a feature m o r e closely resembling anatomically m o d e r n Homo sapiens t h a n the Neandertals, a l t h o u g h the feature is shared b y b o t h specimens and m o r e o v e r it is not shared by all Neandertals. However, c o m b i n e d with other aspects of the occiput such as the form of the occipito-mastoid torus, the lack of relative occipital plane lengthening a p p r o a c h i n g that of later Homo sapiens samples (including both Neandertals a n d more anatomically m o d e r n samples), the separation of inion a n d endinion in Swanscombe, a n d the closed occipital angle (relative to later hominids), the region can h a r d l y be regarded as especially modern. W i t h respect to vault thickness, b o t h specimens are reduced c o m p a r e d with the Homo erectus condition a n d generally resemble the Neandertals in this aspect (Table 2). Steinheim is s o m e w h a t thinner t h a n Swanscombe, especially in the parietal thicknesses. This makes it the more m o d e r n of the two; c o n t r a r y to Vallois, vault thinning is a progressive evolutionary trend. While Steinheim falls within the N e a n d e r t a l range (its thinness is m a t c h e d by the Mt. Circeo and L a Q u i n a 5 dimensions), it is Swanscombe (the presumed "presapiens") that falls above the Neandertal range a n d within the Choukoutien range for several parietal thickness dimensions. Table 3

External parietal angles, calculated by a variation of Delattre and F~nart's procedure; the external rather than internal parietal corners are used. Choukoutien data are from W e l d e n r e i c h (1943) a n d t h e r e m a i n i n g s p e c i m e n s w e r e m e a sured by the author. See text for further details

Sagittal Choukoutien (av.) 3

Temporal Coronal Lambdoidal

91

102

85

82

12 Petralona Swanscombe Steinheim

97 88 87 104 107 101

102 106 97 99 79 84

83 85 88 79 85 88

78 81 88 78 88 87

Cova Negra La Chaise I Lazeret

106 88 115

69 95

88 87 65

97 90

Neandertals (av.) Krapina 16 Saecopastore 1 Spy 2 La Ferrassie 1 Mt. Circeo Le Moustler La Chapelle Gibraltar Mlade~ 1 5 "Modern" average (according to Piveteau)

107

79

92

82

96 115 107 105 114 104 119 99 110 107

90 78 76 76 73 80 67 93 80 78

90 86 97 98 91 103 85 88 90 94

84 81 80 82 82 73 89 80 80 81

107

90

90

73

11

344

M.H. WOLPOFF

Another more modern feature of Steinheim is the lack of separation between the inion position on the weakly developed muchal torus and endinion. This contrasts with all of the Middle Pleistocene hominids (Table 6). A procedure for systematically describing parietal form was introduced by Delattre & F6nart (1958), and utilized by Piveteau in his analysis of the Lazeret and La Chaise parietals (1967, 1970). The procedure begins by defining the circumscribed circle passing through the four points defined by the parietal corners (bregma, pterion, asterion and lambda). The angles defined by the radii from the center of the circle to each of these points can be used to describe parietal form since the longer the side, the higher the angle will be. According to Piveteau's interpretation of Vallois' presapiens criteria, the Neandertal (and pre-Neandertal) forms are expected to have a short coronal border (or a small angle defined between the radii to bregma and pterion) and an elongated temporal border. The former corresponds to less frontal expansion and the latter to the marked cranial length of Neandertals, combined with reduced height. Table 3 shows that the parietal angles for Swanscombe and Steinheim are virtually identical. How these fit in a phylogenetic interpretation of parietal form is obscured by the fact that the result of applying this technique to a large Neandertal sample shows that Neandertal parietal form does not generally fit the descriptions made for Neandertals by the above authors. Moreover, there are no significant differences between the average for Neandertals and the two early anatomically modern Homo sapiens crania from Mlade~. Compared with the data Piveteau presents to represent " m o d e r n " humans, Neandertals (and Mlade~) have shorter temporal borders and longer occipital borders (probably corresponding to their expanded occiputs). It is the Homo erectus sample from Choukoutien that fits the description for Neandertal parietal form ! In Steinheim and Swanscombe, the temporal border is longer than in the Neandertals, and the coronal and sagittal borders are shorter. In all three cases the values lie between the Choukoutien and Neandertal averages. The lambdoidal border is somewhat elongated compared with these samples, but this is hardly a more modern feature since the " m o d e r n h u m a n " value is even shorter yet. In sum, Swanscombe and Steinheim parietal form is remarkably similar and generally midway between the late Homo erectus sample and the Neandertals. While the frontal and face of Swanscombe are missing, this regions is known for Steinheim where an examination of the remaining features reveals a suite of details remarkably like Homo erectus. The main exception is in the height of the face. While Steinheim facial height is usually considered small, a direct measure is not possible because the entire maxilla from the lower nasal border is broken away. Estimates must depend on the degree of alveolar prognathism which is unknown. I have found that a similar measure, from the lower rim of the superior orbital border at midorbit to the toothrow in the molar region correlates highly with upper facial height. This measure is smaller in Steinheim than in the known Neandertal females (Table 4), although the difference is not great. Moreover, any attempt to take the small size of the vault into account results in a relatively large face for Steinheim. Upper facial breadths are also fairly narrow compared with most Neandertal females. However, the lower face and maxilla are generally broader in Steinheim, probably reflecting the development of the masseter muscle. Similarly, the length of the zygomaxillary suture is long compared with the later females. Steinheim facial length is not expanded in response to the marked mid-facial prognathism of the Neandertals (indeed, the value for all five Neandertal females exceeds the facial length of the Petralona cranium).

MIDDLE PLEISTOCENE EUROPEAN

Table 4

345

HOMINIDS

Facial dimensions of Stelnhelm compared with Petralona a n d w i t h the f a c e s o f f e m a l e N e a n d e r t a l s . All m e a s u r e m e n t s (in m m ) t a k e n b y the a u t h o r o n t h e o r i g i n a l s p e c i m e n s

Female Neandertals Gibraltar Saccopastore Tabun Petralona Steinheim Midpoint on inferior orbital border to toothrow Orbito-alveolar height Orbit height Orbit breadth Nasal breadth fmt-fmt fmo-fmo Bijugal breadth Maxillary breadth Facial length (au-fmo) Zygomatic arch length (au-ju) Zygomaxillary suture length Projective zygomatic bone height Supraorbital midorbit height Supraorbital torus length (after Weidenreich)

1

1

83.4

96-6 59.0 36-3 46.5 31.0 133.0 122.0 141-0 112.0 78.4 58.9 44-4

80.6 48-8 32-1 38.8 31.6 108.0 100-4 112.0 107.5 69.6 60.5 33.7

86-0 48.0 39"3 42.8 34"1 114.8 105-5 115-0 102.4 79.8 63.9 27.2

89.0 53.8 38.0 42 '3 32"9 118'0 112-0

36.5 37" 1 36.0 115"5 104.2

94.5 76.6

84"3

32"9 19"8

24-9 16.2

18.0 14.0

27-5 11.4

22-5

20.1

18.9

KrapinaLaQuina C

39.7 40.9

36"7

118.6 108.4 78"9 61-0 30.1

106"5 97.5 102.7 85"0 80-3 65"5 29.6

12.4

24.5 12-3

24"0 14-9

18-9

19.9

24.5

33.5

T h e l e n g t h o f the z y g o m a t i c a r c h is a l m o s t as long as the N e a n d e r t a l female v a l u e , in S t e i n h e i m , a n d c o n s e q u e n t l y the S t e i n h e i m face is flatter transversely. T h e s u p r a o r b i t a l d i m e n s i o n s o f S t e i n h e i m a r e m a r k e d l y g r e a t e r t h a n for almosL e v e r y N e a n d e r t a l female. I n fact, the thickness o f the S t e i n h e i m s u p r a o r b i t a l torus is g r e a t e r t h a n for a n y N e a n d e r t a l male. T h i s is a r e m a r k a b l y a r c h a i c f e a t u r e for so s m a l l a face. V i e w e d in the E u r o p e a n e v o l u t i o n a r y contest, these specimens h a v e a l w a y s p r e s e n t e d a n e n i g m a . W h e t h e r c o m p a r e d w i t h e a r l i e r Homo erectus (from o t h e r areas) or the succeeding E u r o p e a n N e a n d e r t a l s , the c r a n i a l vaults a p p e a r e d to be r e m a r k a b l y gracile. T h i s was especially n o t a b l e in the r e d u c e d structures a s s o c i a t e d w i t h o c c i p i t a l b u t t r e s s i n g a n d the r e d u c e d p n e u m a t i z a t i o n in the t e m p o r o m a s t o i d a r e a (of S t e i n h e i m ) . M o r e o v e r , the small S t e i n h e i m face seemed to c o n f i r m the s m a l l face of the C h o u k o u t i e n 11 rec o n s t r u c t i o n (when it was the o n l y face k n o w n for Homo erectus) a n d t h r e w large faces o f E u r o p e a n N e a n d e r t a l s such as L a Ferrassie a n d L a C h a p e l l e into contrast. Discoveries of the p a s t two decades h a v e p r o v i d e d the basis for a r a t h e r different i n t e r p r e t a t i o n of these specimens. A m o r e a c c u r a t e i d e a o f facial size in Homo erectus has r e s u l t e d from discoveries in I n d o n e s i a ( S a n g i r a n 17) a n d E a s t A f r i c a ( E R 3733). T h e w i d e s p r e a d l a t t e r M i d d l e Pleistocene a p p e a r a n c e of specimens w i t h a g r a c i l i z e d o c c i p i t a l region was c o n f i r m e d w i t h the N d u t u (Clarke, 1976) a n d Sal~ ( J a e g e r , 1975) c r a n i a from Africa. F i n a l l y , in E u r o p e itself n e w discoveries h a v e p r o v i d e d a m o r e a c c u r a t e p i c t u r e o f h o m i n i d v a r i a t i o n d u r i n g this t i m e s p a n . 3. P e t r a l o n a

5

and Bilzingsleben

A l t h o u g h the P e t r a l o n a c r a n i u m was d i s c o v e r e d n e a r T h e s s a l o n i k i in 1959, a n n o u n c e m e n t s a n d b r i e f descriptions d i d n o t a p p e a r u n t i l 1964 ( K a n e l l i s & Savas, 1964) a n d a p a r t from some basic metrics p u b l i s h e d b y several a u t h o r s (Bostanci, 1964; P o u l a i n o s ,

346

~. H. WOLPOFF

1971), the specimen has not yet been fully cleaned or described. The circumstances surrounding the discovery are unclear (there are various differing renditions), and exactly what was discovered is ambiguous. Claims range from the cranium alone to an entire post-cranial skeleton (now missing). The provenience of the specimen is also uncertain, or at least subject to different interpretations, since the individual may have been a burial, may have been lying on a stone slab, or may have been embedded in matrix. Finally, the initial interpretation of the cave fauna as Wiirm in age (Bostanci, 1964; Poulianos, 1967) mislead a number of workers (including this author) to considering the poorly pictured individual as a Neandertal variant. It has become clear that at least part of the cave sequence is considerably older. An analysis of the fauna in the upper part of the cave (a sequence of strata capped by the level from which the cranium is said to have come) suggests a Middle Pleistocene age and ecological conditions indicating interglacial conditions (Kretzoi, 1977). An age equivalent to Steinheim and Swanscombe is not unlikely, although this evidence has also been interpreted to suggest a pre-Mindel age. The circumstances of discovery and the date of Bilzingsleben are far less ambiguous. The site is north of Jena in the DDR. Excavations by D. Mania have resulted in the recovery of two frontal fragments, most of an occiput, two parietal fragments, and an upper molar probably belonging to a single individual (Vl~ek & Mania, 1977). Faunal and geological analysis suggest a Holstein age, once again equivalent to Steinheim and Swanscombe (Mania & Vl~ek, 1977). Taxonomic interpretations of Petralona have ranged from Neandertal to Homo erectus, and its similarities to the Broken Hill remains were noted by a number of authors. Kurtdn (1972) and more recently Stringer (1974), suggested that the specimen should be considered an early form of Homo sapiens. In a multivariate analysis Stringer demonstrated numerous distinctions from Neandertals, and also found the specimen to be unlike Steinheim. Less variety characterizes the taxonomic descriptions of Blizingsleben. The cranium was assigned to Homo erectus by Mania. I was recently able to examine both the Petralona and Bilzingsleben remains. Petralona is a massively buttressed cranium, retaining much of the shape and proportion of Homo erectus males (Plate 1). Vault size, however, is larger than for specimens generally included in the taxon (see Tables 1, 3, 5-8). Moreover, although the vault is extraordinarily broad, the amount of basal pneumatization appears to be reduced compared with earlier specimens. For instance, the nuchal torus is vertically large but does not project prominently. Above it, the occipital plane is fairly filled out. A broad area bulges convexly from a position just superior to the shallow sulcus defining the top of the torus to lambda. Moreover, the cranial contour as seen posteriorally does not attain the normal "gabled" form of Homo erectus. The exact contour is complicated by the marked asymmetry of the vault. The right parietal is close to being vertical while on the left a gentle inferior-lateral curvature begins at about the position of the temporal line. This asymmetry extends on to the occipital and effects the nuchal line which travels horizontally to the left of the midline but curves superiorally and then inferiorally to the right. A possibly related feature is the marked lateral flange of the right temporal posterior border, maximally projecting some 11-7 m m from the vault wall (Plate 2). Unlike the Choukoutien and most other Homo erectus samples, the occipital plane dominates the nuchal plane in length. An inion-like projection appears some 14.5 m m inferior to the midpoint on the nuchal torus. Unfortunately, pneumatization in the mastoid

M I D D L E P L E I S T O C E N E E U R O P E A N HOMINIDS

347

region, and the size of the mastoids themselves, cannot be ascertained because of the yet uncleaned incrustation. Reduced buttressing also characterizes the sagittal torus. The structure is weakly developed, beginning some 25 mm posterior to bregma and terminating well anterior to lambda. There is no corresponding torus on the frontal. The apparent bulge on the sagittal plane is probably better described as an elongated boss. In contrast, the supraorbitals are massive structures. Like Broken Hill and Steinheim, they follow the orbital contours, dipping inferiorally in the region of glabella where, unlike Broken Hill, a slight sulcus extending to nasion separates them. The torus is fairly even in thickness from the medial portion to the approximate midorbit position, where it begins to thin significantly. The face is large by any standard. While there is superficial resemblance to Broken Hill, a number of contrasts occur which may be of significance. The maxilla is p u f e d and the maxillary sinus expanded in a manner clearly foreshadowing the Neandertal condition. However, measures of facial length and midfacial projection are even less than in the Neandertal females (Table 4). The region lateral to the nasal border is convex and there is no canine fossa. T h e nasal spine in Petralona is more projecting than in Broken Hill, and the rim of the nasal aperture coincides with the edge of the nasal floor whereas in Broken Hill the rim extends inferiorally below the floor edge. T h e midface of Petralona projects more anteriorally while below the nose the maxilla drops vertically. In Broken Hill there is more prognathism below the nose. The base of the zygomatic is much narrower in Petralona, and the lack of a palatine torus contrasts with the double ridged torus found in the African specimen. Many of these features distinguish living Europeans from living Africans, and more distinguish Neandertals from their African contemporaries. In my opinion, a sufficient number of metric and morphological features suggest that Petralona cannot comfortably be included in Homo erectus as represented in Asia and Africa. I have no disagreement with suggestions that the specimen be regarded as an early representative of Homo sapiens (Stringer, 1974; Kurt6n, 1972), if not as a specific precursor of Neandertals (Jelinek, 1969). I also regard Broken Hill, Bodo, Ndutu, Saldanha, SaM and the Ngandong sample as early (or archaic) Homo sapiens (Wolpoff, 1980), and I believe that significant grade characteristics can be shown to relate these geographically dispersed crania. In Petralona, these features include the size of the vault, the lack of nuchal torus projection, the expansion of the occipital plane at the expanse of the nuchal plane, the reduction in occipital breadth, the reduction in basal pneumatization, and the tendency of the supraorbitals to follow the superior orbital contours and the reduction of this structure over nasion. Some clade features characteristic of Europe are seen in the sagittal torus reduction, the "squared off" appearance of the anterior tooth row of the palate, the form of the orbit, the lack o f alveolar prognathism, and the expression of midfacial prognathism. Yet, the fact remains that Petralona is more like Homo erectus than the other early European specimens discussed, and markedly more like Homo erectus than the Neandertals. Regarding the specimen as late Homo erectus is compatible with taxonomic schemes that also place Arago and the extra-European specimens of comparable grade in this taxon. Conversely, the whole problem of morphological continuity could also lead to a solution in which Homo erectus is sunk to a subspecific taxon (Jelinek, 1978) within Homo sapiens. I f Petralona can be said to resemble Homo erectus, the Bilzingsleben remains are indistinguishable in any meaningful sense (Plate 3). The central supra-orbital region that

348

M.H.%VOLPOFF

is preserved is massively developed with little " d i p p i n g " and no concavity over nasion. Projection of the torus is somewhat greater than in Petralona, although the thickness where comparable is less (Table 5) and more closely approximates Steinheim. A b r o a d sulcus separates the supraorbitals from the frontal squama which, in turn, does not a p p e a r to have a marked vertical angulation. T h e whole region is reminiscent of O H 9, differing from the Choukoutien morphology in torus thickness, lack of a depressed nasal root, and absence of a frontal boss. Table 5

Comparable frontal dimensions for pre-Riss Middle Pleistoc e n e E u r o p e a n h o m i n i d s (in r a m ) . All m e a s u r e m e n t s t a k e n by the author on the original specimens

Bilzingsleben Petralona Steinhelm Midorbit projection of torus (from internal) Medial projection of torus (from internal) Supraorbital torus length (after Weidenreich) Supraorbital thickness : medial midorblt

30.9

20.7

25"0

19"5

24-6

22-5

20-1

18.2 17-1

22-6 19.8

17.8 16.2

The Bilzingsleben occiput is very b r o a d (Table 6). T h e length of the occipital plane is similar to Petralona, and there is a significant separation between inion (not present as a prominence but defined on the torus midline) a n d endinion. Unlike Swanscombe, there is no Neandertal-like depressed area at the midline just above the nuchal torus. Instead, the very shallow sulcus is discontinuous at the midline where an almost straight superior extension of bone only begins to curve anteriorally just inferior to lambda. I n contrast, the chord/arc index shows that the parietal border of the occuput is strongly curved. Cranial vault thickness measures for Bilzingsleben cannot be compared with Petralona. However, the massiveness of the bone is indicated by the fact that thickness at b r e g m a a n d at inion exceed the Choukoutien average [although they fall within the observed range (Table 2)]. 4. O t h e r S p e c i m e n s

Adult European cranial remains dated to the middle and later portions of the Middle Pleistocene include Vdrtessz6116s which m a y be earlier than the specimens discussed above, and remains from Arago, Biache and La Chaise which m a y be later. While I have been fortunate in being able to examine all of these, a complete discussion of Biache a n d the Arago remains must await their detailed publication. T h e occipital from the Mindel dated site of V6rtessz6116s (Kretzoi & Vertes, 1965) has been discussed as a representative of late Homo erectus (Wolpoff, 1971, 1977) or Homo erectus or sapiens (Thoma, 1966, 1969). Von Koenigswald noted its similarities to Petralona (1967). Some of the differences from Homo erectus parallel those of Petralona. These include the size of the occiput and the high occipital angle. Affinities to Homo grectus as represented in other areas are suggested by the large nuchal plane relative to the occipital plane (Table 7), the magnitude of the distance between inion a n d endinion

MIDDLE

Table 6

PLEISTOCENE

EUROPEAN

349

HOMINIDS

Comparable occipital dimensions for pre-Riss Middle Pleistocene E u r o p e a n h o m i n i d s (in m m , s e e a l s o T a b l e 1). A l l measurements taken by the author on the original specimens

Bilzingsleben Petralona V6rtessz6116s Steinheim

Swanscombe

Biasterionic breadth

138.6

119.0

128.3

102.5

121.8

Lambda-asterion arc chord index

103.0 88.3

97.0 87-3

101.0 90"3

101-0 86.7

104 91.6

116.6

111.1

111.8

116.5

113.5

54.0 50.5

68.0 63.6

53.0 49.8

62.5 60.4

63.0 58.0

106.9

106.9

106.4

103.8

108"6

Lambda-inion arc chord index Distance of inion above endinlon

24.5

27"8

0.0

15.8

Bone thickness: Occipital-mastoid suture (near asterion) Larnbda Inion Endinion

16.5 10"9 17.0 13.5

14.3 10.0 16.0 I0.0

7"5 8.5 11.1 11.1

11.8 10.4 9.0 14.5

(Table 6), the retention of a sulcus inferior to the nuchal torus, a n d the thickness of the occipital at asterion and inion. I n these (and other features) Vdrtessz6116s is more like Homo erectus than Petralona is, but the general resemblance between these specimens is marked. Moreover, the few comparisons possible with the more fragmentary Bilzingsleben occiput (not including the relative size of the nuchal plane which cannot be determined for the G e r m a n specimen) suggest that Bilzingsleben is even more Homo erectus-like. The lambda-inion distance is slightly less while the biasterionic breadth is m a r k e d l y greater (Table 6). T h e newly discovered Biache cranium (Vandermeersch, 1978) is dated to an interstadial within the Riss. While only the occiput, temporals a n d parietals remain of the vault (a palate and some post-canine teeth were also discovered), the resemblance to Swanscombe is extraordinary in both size, proportions, and morphological detail. Biache could be described as Swanscombe with a little more occipital/parietal flattening in the lambdoidal region. The mastoids (not preserved in Swanscombe) are virtually identical to Steinheim in size. T h e nuchal torus is very poorly expressed, a n d retains neither an inferior nor a superior sulcus. In some features the cranium is even less like Homo erectus than Swanscombe. T h e occipital plane is larger relative to the nuchal plane, and vault thickness is less. As seen from the rear, m a x i m u m parietal breadth is about midway up the vault, and the general contour is semicircular. A similarly gracile occipital was recovered at La Chaise (Piveteau, 1970). Morphologically La Chaise 2 combines a reduced expression of the nuchal torus with a high occipital angle. The occipital plane is highly curved (Krukoff, 1970), and has a "filled out" appearance. In absolute dimensions it is a small specimen, exceeding only Steinheim in o c c i p i ~ l breadth. T h e occipital plane is barely larger than the nuchal plane (art archaic feature). A temporal from another individual, La Chaise 3, resembles Biache in its small mastoid

350

M. H. WOLPOFF

Table 7

Saglttal d i m e n s i o n s o f the occiput and the occipital index for Middle Pleistocene Europeans, and Neandertals w i t h both nuchal and occipital planes preserved

Nuchal Plane Length

Occipital Plane Length

Index

Petralona Vrrtessz6116s Swanscombe La Chaise 2

60-6 57.0 51.0 47-9

63.3 49-8 58.0 50-2

95.3 114-5 87.9 95.4

Gibraltar Saccopostore 1 La Ferrassie 1 La Chapelle Ehringsdorf H

47.7 47.0 45-5 39.9 46.9

56.0 56.8 62.3 59.6 57.5

85"2 82.7 72-0 66.9 81.6

process and short temporal squama. La Chaise 1 is the most complete calotte [the specimen announced by Piveteau (1972) as lacking a supraorbital torus has been discovered to be Mesolithic in age]. Although missing the anterior portion of the frontal, its marked parietal breadth and the semi-circular shape of the vault seen from norma occipitalis is an important similarity to Biache since it foreshadows the usual Neandertal condition. However the vault is small (estimated capacity is 1065 ec), low and thick. Moreover, the parietal angles (Table 3) are more like Homo erectus than any other Middle Pleistocene European crania, combining sagittal shortening and temporal elongation. Dated to the Riss I I I interstadial, the La Chaise remains are probably the latest of those discussed here. The only Riss site with facial material is Arago (de Lumley & de Lumley, 1974). T h e Arago 21 cranium retains the face and frontal. Although both are fairly distorted, numerous morphological features and some metrics can be accurately ascertained. T h e face is intermediate between Steinheim and Petralona in size (de Lumley, 1976). Facial heights closely approach Steinheim, breadths of the upper face are more like Petralona, while middle and lower facial breadths are midway between the two or again more similar to Steinheim. The few ascertainable dimensions of the frontal are almost identical to Petralona. The supraorbitals are fairly thick, following the orbital contours and separated from the frontal squama by a broad suleus. T h e y differ somewhat from the tori discussed above, although like Steinheim and Petralona a broad depression over nasion separates the right and left elements. A weakly developed incisura frontale forms the medial inferior base of a broad groove that separates the supraorbitals into medial and lateral elements. The medial element is much thicker than the lateral, and exceeds both Steinheim and Bilzingsleben. Midorbit thickness, measured on the lateral element, is less than Steinheim while the most lateral supraorbital thickness is somewhat greater. Supraorbital length, as defined by Weidenreieh, is about the same as Steinheim. The right orbit and the left infraorbital region show the least distortion and crushing. T h e orbit is relatively low and broad, and the lateral border is very thick. T h e zygomatie process of the maxilla is convex, lacking a canine fossa, and the maxillary puffing characteristic of the later Neandertals is well developed. T h e whole midfaee angles sharply posteriorally (the apparent facial flatness on the right side is an artifact of crushing), and the lower border of the zygomatic process angles smoothly into the alveolar region of the bone. The resemblance to the morphology of the Saccopastore 2 face

MIDDLE PLEISTOCENE EUROPEAN HOMINIDS

351

(Sergi, 1948) is striking, although the Italian Neandertal lacks the marked supraorbital thickening and has a considerably taller face. While the dentition is usually regarded as large, comparisons to Bilzingsleben and Petralona (possible for the M 1 only) suggest some reduction. The summed area of the three molars is almost exactly midway between Petralona and Steinheim, and the Arago 21 M 3 shows the greatest amount of relative reduction. The Arago 21 maxillary dentition is not the largest at the site. A comparison of posterior tooth areas with the composite dentition of Arago 1 (W, M x and M S) shows Arago 21 to be only 80% as large. Moreover, the Arago 1 mandibular dentition has only 9 0 % the size of the Arago 13 mandible for comparable teeth (P4, Ms). Since the Arago 1 maxilla has larger teeth than Arago 21 while the Arago 1 mandible has smaller teeth than Arago 13, it is unlikely that the Arago 21 maxilla and the Arago 13 mandible make a good morphological "fit", or that Arago 13 could be expected to resemble the Arago 21 mandible. 5. D i s c u s s i o n

Variation in the Middle Pleistocene European hominids probably has three main sources: time, sex and idiosyncrasy. Given the size of the sample, the problematic dating for many of the individuals, and the lack of large contemporary samples from other regions, it is unlikely that the exact contribution of each will ever be fully clarified. If all the dates normally proposed are accepted, it would appear that individuals more like Homo erectus overlap in time with individuals usually regarded as representing early Homo sapiens. However, this need not be the case. Recent papers have suggested that both Steinheim and Swanscombe may be later than is generally proposed. Bordes (1968) has suggested a Riss date for both of these. If Steinheim were actually Riss in age, and Swanscombe Riss or Riss/Wfirm, then all of the more archaic appearing specimens (V6rtessz6116s, Petralona and Bilzingsleben) would be earlier in time. While this need not contradict the discussion below, it complicates any determination of sex-based variation by introducing the additional contribution of differing expressions of evolutionary trends. Apart from the dating problem, much of the variation in this sample could be interpreted as the result of marked sexual dimorphism. The robust group, presumably the males, have a greater expression of Homo erectus-like features. Nonetheless, in the discussion of Steinheim and Swanscombe it was pointed out that where these specimens differ from Neandertals, it was in the direction of Homo erectus in virtually every case. Parietal form, proportions of the occiput, vault thickness, development of the supraorbital torus (length, breadth and evenness of thickness over the orbit), narrowness of the upper face, and the size of the zygomatic process of the maxilla all point in this direction. While Stringer's discriminate function clearly separates the Steinheim and Petralona faces (1974), the biological basis for this separation could as well be sexual as phylogenetic. The major differences between the gracile and robust samples are expressed in the size and thickness of the vault (Table 1, 3, 5-8) reduction of the occipital and sagittal buttresses, the development of the muscle markings, the curvature and height of the forehead, size of the supraorbitals, the prominence of the glabellar region, the size of the face (Table 6), the development of the zygoma and the zygomatic process of the maxilla, nasal breadth (narrower in Petralona), the orbital index, and the size of the post-canine

352

M.H. WOLPOFF

Table 8

Comparison of vault dimensions and saglttal projections from t h e a u r i c u l a r p o i n t f o r S t e i n h e i m a n d P e t r a l o n a (in r a m ) . All m e a s u r e m e n t s w e r e t a k e n b y the a u t h o r o n o r i g i n a l s p e c i m e n s . See T a b l e s 1, 3 a n d 5-7 f o r a d d i t i o n a l v a u l t comparisons

Petralona Steinheim Cranial length Asterion-glabella Basion-nasion Glabella-lambda Cranial breadth Biparietal breadth Minimum frontal breadth Maximum frontal breadth Biauricular breadth Breadth across mandibular-fossae Nasion-bregma Glabella-bregma Breadth/Length index Height/Length index Parietal]Cranlal breadth index Auricular projections (midline) to: nasion glabella sagittale bregma lambda opistocranion inion basion

210.0 179"0 111.5 188"8 168.0 150.0 111.8 120-5 t54"8 146.5 107.9 104.8

179.0 148.5 94"8 170.0 130.0 126-8 95"5 111.5 113-5 112-7 99.2 84.5

80.0 60"7 89.3

72.6 58"3 97.5

98.9 109.0 106.9 103.4 102.2 100-1 98.0 12.5

89"8 100.1 96-0 97.5 105.6 94.8 90.0 6"3

teeth. The fact is that this list closely corresponds to the normal cranial features that are used in sex determination for living humans (Krogman, 1962). This correspondence is probably not accidental. In any event, the parallel (although not identical) expression of features resembling Homo erectus in the more gracile and more robust samples, a n d the fact that these differ from each other in a suite of features that distinguish males a n d females in modern Homo sapiens makes sexual dimorphism a suspected important contributing factor to the variation observed. A specific comparison of Petralona and Steinheim (Table 9) suggests a truly marked degree of dimorphism when compared with m e a n values for living populations. However, average dimorphism for the European Neandertals comes close to matching the marked expression of dimorphism in the Petralona/Steinheim comparison. Moreover, comparison of the two most complete male and female Neandertal specimens (La Ferrassie and Gibraltar) comes closer yet. Nevertheless, in virtually every case the Neandertal dimorphism (either comparison) is less than the Petralona/Steinheim comparison although the differences are not great and the Neandertals are m u c h more like these earlier specimens than like living humans with regard to their expression of dimorphism. T h e sample size is too small to determine whether the slight reduction between the values suggested for Middle Pleistocene European hominids and the later Neandertals represent an evolutionary trend. I f there was such a trend, the a m o u n t of reduction involved is very small. The indication of marked dimorphism in the cranial a n d facial

MIDDLE

Table 9

PLEISTOCENE

EUROPEAN

Comparison of percent sexual dlmorphlsm (male divided by f e m a l e ) i n P e t r a l o n a a n d Stelnhelm~ m a l e a n d f e m a l e N e a n d e r t a l a v e r a g e s , La F e r r a s s i e a n d G i b r a l t a r , a n d a m e a n v a l u e for l i v i n g p o p u l a t l o n a l a v e r a g e s

Neandertal*

Modernt Male/Female

A

t

Male/Female

La Ferrassie/ Gibraltar

117 129 108 120 118

110 109 105 116 115 112

116 109 99 121 119 118

105 103 104 104 105 107

120 121 123 113 98

112 116 106 120 98

114 124 106 109 99

109 105 106 104

Petralona[ Steinheim Cranial length Maximum cranial breadth Maximum frontal breadth Cranial height Nasion-basion Upper facial height Inferior upper orbit border to tooth row Orbito-alveolar height Outer orbital breadth (fmt-fmt) Midfacial breadth (zm-zm) Nasal breadth

353

HOMINIDS

* Average figures: females are Gibraltar, Saccopastore 1, Krapina C, La Ouina 5, and Tabun; males are La Chapelle, La Ferrassie, Saccopastore 2, Mt. Circeo. The specimens were chosen for facial completeness. I" Average of populational means from Howells (1973). remains of Neandertals is of interest insofar as it contrasts with the reduced degree of dimorphism in the size and morphology of the postcranial remains (Heim, 1974, 1976; Trinkaus, this volume). Finally, that the extent of dimorphism indicated by the Petralona/Steinheim comparison is not unusual in a Middle Pleistocene context is suggested by comparisons of roughly contemporary African cranial remains. I f Broken Hill, Saldanha and Bodo are considered males, and Sal6 and Ndutu are considered females, the expression of dimorphism and its pattern of variation are quite similar to Europe. The characteristics that suggest Steinheim and Swanscombe are females, and Petralona, Bilzingsleben and V6rtessz6116s are males may also be used to indicate sex for the other specimens discussed. O n the basis of their small size, weakness of the nuchal torus, gracility of muscle attachments, and small mastoids, all three of the La Chaise cranial fragments would appear to be female. Similarity between Biache and Swanscombe indicates that this specimen could also be regarded as female. The Arago 21 face and anterior vault is more problematic since the size of the supraorbitals and m a n y of the robustly developed facial features led to the initial assumption that the specimen was male. I believe that several factors combine to suggest that the opposite may be the case. To begin with, the Arago 21 dentition is smaller than the only other maxillary tooth set from the site, Arago I, and the Arago 1 mandibular teeth are in turn smaller than the teeth of the male mandible Arago 13. Thus, tooth size would not obviously indicate that the specimen was male. While the size of the supraorbitals are indeed large, the most dramatic contrast is with the Neandertals. The Arago 21 supraorbitals are smaller than those of Petralona and Bilzingsleben, and only exceed Steinheim in some dimensions. Large supraorbitals, in other words, may be more of a reflection of the date

354

M.H. WOLPOFF

of the specimen than of its sex. Moreover, the size of the supraorbital torus is not a good sex discriminator for the Neandertal faces of known sex. T h e best sex indicators for Arago 21 are found in the face. Sexual dimorphism in Neandertal faces is most pronounced in the facial height measures, and midfacial breadth (Table 9). These are the measures in which Arago 21 most closely resembles Steinheim (near identity in most cases). Neandertal sexual dimorphism is least pronounced in the breadths of the frontal, the outer orbit, and the nose. These are the measures in which Arago 21 lies between Petralona and Steinheim. In sum, I believe it likely that Arago 21 is a female. Considered as a female, Arago 21 can be seen to differ from the Neandertal females (Tables 6, 9). These later specimens developed a suite of facial features which are probably associated with their expanded maxillary incisors but generally reduced masticatory apparatus. For instance, upper facial height in the Neandertal females is somewhat greater although orbitoalveolar height is reduced in spite of the fact that orbit height is expanded. T h e change in facial height is mainly in the lower (incisor bearing) maxilla. Similarly, expansion in the breadth of this region is indicated by greater bicanine a n d nasal breadths in the Neandertal females. Masticatory function related decreases include the size of the zygomatic process of the maxilla a n d the decrease in maxillary (midfacial) breadth. Supraorbital reduction is probably also related to decreased masticatory-related stress (Wolpoff, 1980), although an additional contributing factor was the increased ability of the higher and more bulging Neandertal female foreheads to resist bending resulting from incisor loading (Endo, 1966, 1970). A more direct measure of decreased masticatory stress is found in the marked posterior tooth size reduction (Wolpoff, 1978). I f sex determination as suggested here is correct, to what extent is the dimorphism estimate confused by the earlier date for most of the males? Evelutionary trends within the Middle Pleistocene European sample cannot be ascertained. This is because there are no reasonably complete cranial remains of later males to compare the earlier male sample with, and the uncertainties in dates for Steinheim a n d Swanscombe make it problematic to seek trends between these specimens and the Riss females. I f the evolutionary trends that took place during this period involved the "gracilization" of both sexes, it is possible that the extent of dimorphism suggested here is partially an artifact of phyletic evolution. T h e probability is that this would tend to result in an overestimation of dimorphism. Yet, comparison with the Neandertals indicates that the extent of overestimation could not be great unless there was a trend to increase the a m o u n t of dimorphism between the Middle and U p p e r Pleistocene samples, a n d based on all of the evidence available this is unlikely. I believe the estimate of Middle Pleistocene sexual dimorphism under the assumption that the specimens are correctly sexed is probably not significantly inaccurate even if this estimate includes some a m o u n t of variation due to phyletic evolution. Moreover, this estimation is not unlike roughly contemporary dimorphism in Africa, or later dimorphism in the Neandertals. While none of this can prove that sexual dimorphism is a valid explanation for the observed variation in the European Middle Pleistocene hominids, I believe it is the most credible explanatory hypothesis. T w o alternative approaches could be used in the interpretation of this sample. T h e first of these is the multivariate approach, probably best expressed in a recent p a p e r by Stringer (1978). Cranial angles, indexes a n d the Penrose size and shape statistic are used to compare the more complete of these specimens to each other a n d to a wide range

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of later hominids. While many of the conclusions parallel those suggested here (for instance, the lack of facial projection in Steinheim and Petralona as compared with the later Neandertals), these are burried in a series of suggestions that are so disparate that they defy summary. A Penrose shape analysis using the Arago 21 face as a reference specimen places Steinheim further from it than Neandertal faces, Petralona or Broken Hill. Petralona, in turn, is said to most closely resemble Choukoutien, Broken Hill, Jebel Irhoud and the Neandertals in an analysis of vault measurements. Yet, in the analysis of facial angles, Steinheim, Arago 21 and Petralona (along with the Saccopastore crania) are said to form a distinctive group unlike the Neandertals. In all, there is nothing in Stringer's analysis that detracts from previous conclusions regarding the usefulness of multivariate statistics for providing insight into paleoanthropological problems (Wolpoff, 1976). A second alternative interpretation of these data could result from a strict application of the cladistic approach (cf. Tattersall & Eldredge, 1977; Delson, 1977). Comparisons of shared derived features in this sample would almost certainly align the (proposed) male sample with Homo erectus, and result in the contention that two hominid lineages as represented by distinct morphotypes shared Europe in the Middle Pleistocene. While such a contention cannot be completely dismissed or unambiguously disproved, it so greatly contradicts the shared evolutionary trends of the two (i.e. " m a l e " and "female") samples, ignores the sexually diagnostic differences between them, and flies so strongly in the face of experience with normal human variation, that it does not appear to be a viable alternative hypothesis. If neither a multivariate nor a straightforward cladistic approach is used, there remains the problem of how to treat this European sample taxonomically. A number of factors combine to make the taxonomy of this sample problematic. It is a sample and not a population. The sample represents a broad timespan but not every specimen can be accurately dated within it (either absolutely or relative to other specimens). There appears to be a marked expression of sexual dimorphism. Finally, a good case can be made for evolutionary continuity between earlier Homo erectus samples outside Europe and later Homo sapiens samples living in the same areas. When only Steinheim and Swanscombe were known, paleoanthropologists benefited from the "natural gap" in the fossil record that clearly delineated these specimens from Homo erectus in time and morphology. The much more complete sample now available seems to raise virtually every problem that can stem from applying a discrete biological classification to an evolutionary continuum (Mayr, 1969). Either the sample represents a new evolutionary species or it can be included within one or more previously defined species. The former is so unlikely that it probably requires no discussion. I f the sample is to be included in previously described species, the alternatives are three: it represents Homo erectus, it represents Homo sapiens, or part of the sample should be allocated to each. In the latter case, a cladistic approach would be the least ambiguous, but it probably would reflect biological reality in the most inaccurate way. An attempt to divide the sample on temporal grounds must resort to arbitrary criteria and moreover is clouded by problems in dating Petralona, if not also Swanscombe and Steinheim. Yet, an attempt to include the entire sample in either species of Homo is equally problematic (Bitsborough, 1976). Including it in Homo erectus ignores the virtual identity of some of the specimens (especially but not exclusively the later ones) with subsequent samples from Europe while including it in Homo sapiens results in the

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equivalent problem of the virtual identity of specimens such as Bilzingsleben and V6rtessz6116s with Homo erectus as represented in other regions. In the view of this author, there is no objective solution to these problems. Any solution is both subjective and arbitrary, and it is probably the case that no one consistent approach is better than any other. Mayr (1969) suggests that in cases such as this the ultimate classification be judged by its reflection of biological reality, its consistency with the treatment of other samples, and the amount of information retained in the classificatory scheme. M y own preference is to retain the earlier taxonomic designations which allocate Steinheim and Swanscombe to Homo sapiens, and suggests using the arbitrary criterion of the end of the Mindel glaciation (or its equivalents) for delineating the Homo erectus/Homo sapiens boundary. This would result in considering V6rtessz6116s as late Homo erectus, Bilzingsleben and most of the other specimens discussed as early Homo sapiens, and Petralona as possibly either depending on its date. Yet, I would prefer not to classify these specimens at all. The statement that this sample represents a segment of a single evolving lineage broadly descendent from earlier Homo erectus populations and broadly ancestral to European Neandertals is a much more accurate assessment of the phylogeny of these early European hominids.

6. Summary Steinheim and Swanscombe are strikingly similar where comparable, although Steinheim is a much smaller cranium. Where these two crania differ from the Neandertals, it is generally in the direction of being more like Homo erectus. Petralona, Bilzingsleben and V6rtessz6116s are markedly larger, more robust, and even more similar to Homo erectus (especially the latter two) than the above. Yet, they differ from Steinheim and Swanscombe mainly in metric and morphological features that distinguish males from females in both living populations and in Neandertals. The most likely hypothesis is that they represent dimorphs in a lineage notably more dimorphic than living people, although not significantly more dimorphic than the Neandertals. Applying the same sex distinguishing criteria to the Riss hominids suggests that all of the best preserved crania remains, including Arago 21, are female. While the phylogeny of the sample seems unambiguous, consideration of its taxonomy brings all of the problems involved in applying Linnean nomenclature to an evolving lineage into focus. This research was supported by NSF grants BNS-76-82729 and BNS-75-21756 and a grant from the National Academy of Sciences, Eastern European Exchange Program. I would like to express my gratitude to the following individuals and institutions for access to study the specimens discussed in this paper, and their help and cooperation during my visit: K. D. Adam, Landesmuseum Stuttgart, Ludwigsburg; H. Bach, Instituts Ptir Anthropologie und Humangenetik, Jena; A. Acanfora, Museo della Preistoria del Lazio, Rome; I. Crnolatac, Ge61osko-paleontolo~ki muzej, Zagreb; H. Delporte, Mus6e Des Antiquit6s Nationales, Yvelines; E. Feustel, Museum fi~r-und-Frtihgeschichte Thtiringens, Weimar; F. Fulep, Magyar Nemzeti Muzeum, Budapest; J. Heim, Musee de l'Homme, Paris; J. Jelinek, Anthropos Institute Moravske Museum Anthropos, Brno; H. Joachim, Rheinisches Landesmuseum, Bonn; A. Leguebe, Institut Royal des Sciences Naturelles, Bruxelles; M.-A. de Lumley, Laboratoire de Pal6ontologie Humaine et de Prfhistoire, Marseille; M. Malez, Istra~iva6ki centar J A Z U , Jedinica za paleon-

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t o l o g i j u i g e o l o g i j u k v a r t a r a , Z a g r e b ; D. M a n i a , L a n d e s m u s e u m H a l l e , H a l l e ; J . Melentis, Institute of Geology and Paleontology, Thessaloniki; J. Piveteau, Laboratorie d e P a l 6 o n t o l o g i e des V e r t 6 b r 6 s et de P a l 6 o n t o l o g i e H u m a i n e U n i v e r s i t y o f Paris, P a r i s ; E. S e r g e N a l d i n i , I n s t i t u t o I t a l i a n o D i P a l e o n t o l o g i a U m a n a , R o m e ; C. S t r i n g e r , D e p a r t m e n t o f A n t h r o p o l o g y British M u s e u m ( N a t u r a l H i s t o r y ) , L o n d o n .

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