Journal of Human Evolution 46 (2004) 105117

Oreopithecus bambolii: an unlikely case of hominidlike grip capability in a Miocene ape

Randall L. Susman*
Department of Anatomical Sciences, School of Medicine, Stony Brook University, Stony Brook, NY 11794-8081, USA Received 15 July 2003; accepted 23 October 2003

Abstract Oreopithecus bambolii, an ape from the late Miocene of Italy, is said to possess a hand capable of a precision grip like that of humans. Relative hand length, proportions of the thumb, and morphological features of the thumb and wrist were adduced to support the idea that Oreopithecus had a hand that closely matched the pattern in Australopithecus. A reappraisal of earlier arguments and comparisons of Oreopithecus with humans, apes, and Old World monkeys, reveals that Oreopithecus had an essentially apelike hand that emphasized apelike power grasping over humanlike precision grasping.  2003 Elsevier Ltd. All rights reserved.
Keywords: Hand morphology; Hominoid evolution; Limb proportions; Miocene ape; Oreopithecus; Precision grasp; Thumb

Introduction Moya ` -Sola ` et al. (1999:313) have suggested that the Miocene ape Oreopithecus bambolii possessed a hand that . closely matches the pattern of early hominids, presumably as a response to similar functional demands. Moya ` -Sola ` et al. concluded this from 1) a novel restoration and reconstruction of the hand of specimen IGF 11778 (Figs. 1 and 2), 2) a restoration and reconstruction of the subadult hand BA #140, and 3) assessment of assorted individual hand bones assigned to Oreopithecus.
* Corresponding author. Tel.: +1-631-444-3125; fax: +1-631-444-3947 E-mail address: randall.susman@stonybrook.edu (R.L. Susman).

The authors determined from their appraisal of thumb length, hand length, and other features of the hand, that Miocene Oreopithecus had a hominidlike precision grip.1 They suggested that humanlike precision grasping evolved rst in Oreopithecus, around 7.5 Ma (Rook et al., 2000), and later, around 3.5 Ma in Australopithecus afarensis. The conclusions reached by Moya ` -Sola ` and colleagues revive the earlier notion of de Terra
1 Napier (1956) described the precision grip and Landsmeer (1962) discussed its anatomical basis. The power grip is one in which the object is held between the palm and the exed ngers, with the thumb supporting the ngers. The power grip is used when the full strength of the hand is needed and precise holding is subordinate. The precision grip dominates when there is a need for ne control. Humans, apes, and monkeys use both types of grips to varying degrees.

0047-2484/04/$ - see front matter  2003 Elsevier Ltd. All rights reserved. doi:10.1016/j.jhevol.2003.10.002

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Fig. 2. Left hand of Oreopithecus (IGF 11778). Note the complete set of 4 middle phalanges and 5 distal phalanges. Of the thumb bones, only a partial distal phalanx remains. Proximal phalanges (pphs) III and IV might be reversed; pph V is the least distorted of the group. The metacarpals are badly deformed and severely crushed from side to side rendering the metacarpal heads uninformative. The scaphoid and lunate remain from the proximal carpal row. The scaphoid is relatively well preserved. The capitate and hamate are present from the distal carpal row and are in poor condition.

Fig. 1. Partial skeleton of Oreopithecus bambolii (IGF 11778) from Baccinello, Italy. The skeleton is badly crushed and most bones are distorted. The femoral head appears intact and indicates a body weight of 32 kg (Jungers, 1987). The limbs suggest that Oreopithecus had orangutanlike proportions (Jungers, 1990). All photographsR. Susman.

(1956) and Hu rzeler (1958, 1960) suggesting hominid morphological anities of Oreopithecus and run counter to the ndings of Straus (1963), Szalay and Delson (1979), Susman (1985), Jungers (1987, 1990), Sarmiento (1987), Sarmiento and Marcus (2000), Harrison (1991), and Harrison and Rook (1997), all of whom view Oreopithecus as ape- or monkeylike.

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Two sets of fossils, BA #140 and IGF 11778, represent partial hands of Oreopithecus. Both sets contain specimens that are broken and plastically deformed (Fig. 2). BA #140 is smaller than IGF 11778 and represents a sub-adult individual as judged by the lack of epiphyseal fusion in some bones. In addition to the above, other bones such as BA #130, a partial distal phalanx, are included in the discussion of hand morphology and function (Moya ` -Sola ` et al., 1999). I had the opportunity to study the IGF 11778 skeleton and other isolated remains of Oreopithecus in Florence in 1974. In 1984 I had an opportunity for further study of Oreopithecus while specimens were on loan to the American Museum of Natural History. My recent requests (in 1999 and again in 2003) for permission to see the fossils were not granted.

Thumb length and morphology in Oreopithecus (IGF 11778 and BA #140) Following Napier (1956), Moya ` -Sola ` et al. (1999: 313) stated Thumb/nger proportions signicantly reect the quality of precision grip capabilities. Based on their reconstruction of the BA #140 and IGF 11778 hands, Moya ` -Sola ` et al. concluded that Oreopithecus had a relatively long, and therefore humanlike, thumb. In their Fig. 2a (Fig. 3 here), they illustrated a regression of the length of the index ngers proximal phalanx on the length of the thumbs proximal phalanx. Their plot of the pollical proximal phalanx of Oreopithecus nds it relatively long (and humanlike) compared to those of apes and most monkeys. The position of BA #140 in Moya ` -Sola ` et al.s Fig. 2a is improbable. To achieve this point on the plot of proximal phalanx I and II lengths, the lengths of both the index and pollical proximal phalanges of BA #140 would have to be essentially equal. Taking the antilogs of x and y values for this point, one arrives at lengths of approximately 20 mm for both the thumb (x) and index (y) phalanges. In addition to Oreopithecus, Moya `Sola ` et al. identied a number of extant primates as having long thumbs. These include humans and papionins (baboons and geladas). However,

according to data on these taxa I collected at the American Museum of Natural History in New York, and data from Etter (1973), thumb proximal phalanges are never as long as their counterparts in the index nger. In humans, with the longest thumb among the primates, the pollical proximal phalanx is only 76% of the length of the index proximal phalanx. The corresponding values for the pollical and indicial proximal phalanges are 73% in geladas, 65% in baboons (P. ursinus), 55% in gorillas, 55% in bonobos, 53% in chimpanzees, and 39% in orangutans (Table 1). I am not aware of any extant anthropoid that has a pollical proximal phalanx and indicial proximal phalanx of equal length, as suggested by Moya ` -Sola ` et al. In addition to Oreopithecus, in Fig. 2a these authors also identied individuals in a mixed group of baboons (Papio) and geladas (Theropithecus) as having pollical and indicial proximal phalanges of equal length. However, a plot of Papio ursinus data (n=5) and data from Etter (1973) on Theropithecus gelada (n=5), Papio hamadryas (n=12), and P. cynocephalus (n=8), indicate that all have a considerable disparity in the lengths of the thumb and index proximal phalanges. My baboon data and those of Etter are plotted, along with data of Moya ` -Sola ` et al., in Fig. 3. Comparison of Fig. 3 with Fig. 2a of Moya ` -Sola ` et al. indicates that they have misassigned and/or mismeasured phalanges of Oreopithecus, as well as those in their comparative sample of monkeys and apes. As in the case of BA #140, a closer look at IGF 11778 indicates a similar problem with the thumb length reconstruction in this specimen. Comparing length proportions of the proposed proximal phalanges of the thumb and index nger of IGF 11778 as plotted in Fig. 2a of Moya ` -Sola ` et al. yields lengths of approximately 32 mm for the purported pollical proximal phalanx (x) and 39 mm for the indicial proximal phalanx (y). This yields percentage value of 82% for the relative length of the pollical proximal phalanx in IGF 11778. This value makes the thumb of IGF 11778 longer than that of humans (Table 1). My own observations of the original IGF 11778 fossils indicates that proximal phalanx V, not II, measures 38.9 mm. It appears that a fth proximal

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Fig. 3. Length of the proximal phalanges (pphs) of the thumb (x) and index nger (y) in selected anthropoid primates, redrawn from Fig. 2a of Moya ` -Sola ` et al. (1999).The positions of Oreopithecus and BA #140 indicate that the thumb and index nger have pphs of equal length. Likewise, Papio and Theropithecus individuals plotted by Moya ` -Sola ` et al. also appear to have thumb and index pphs similar in length. Etters data (*) for Theropithecus gelada, Papio cynocephalus, and P. hamadryas are plotted. Etters data suggest that the papionins, like other anthropoids, including Oreopithecus, do not have thumb and index proximal phalanges of equal length (see text for explanation of these data). Data of Susman(+) concur with those of Etter. The position of IGF 11778 is improbable. The suspect points (BA #140, IGF 11778, Papio, and Theropithecus) are likely based on misidentied phalanges.

phalanx was mistaken for a second proximal phalanx by Moya ` -Sola ` et al. In addition, there appear to be problems with the identication of the pollical proximal phalanx. Together, these problems appear to account for the discrepancies in the data of Moya ` -Sola ` et al. and those presented here (see below). The regressions in Fig. 2a of Moya ` -Sola ` et al. (1999) suggesting that Oreopithecus had a relatively long thumb are suspect. A glance at their Fig. 3 (reproduced as Fig. 4 here), which in part depicts the IGF 11778 hand, reveals that the bone represented by Moya ` -Sola ` et al. as a pollical proxi-

mal phalanx is, in fact, a manual middle phalanx. This is especially evident in Fig. 3b (Fig. 4b here). A few key morphological features reveal the true identity of the bone assigned by Moya ` -Sola ` et al. to the thumb of IGF 11778. Both the body and the basal morphology of the bone seen in Fig. 3b of Moya ` -Sola ` et al. are unlike that of pollical proximal phalanges see (Fig. 5 here). Human pollical proximal phalanges are stouter, with a more hour-glasslike outline. Apes have less robust pollical proximal phalanges with more parallel-sided, less hour-glass shaped bodies. In the middle phalanges of IGF 11778, as in other

R.L. Susman / Journal of Human Evolution 46 (2004) 105117 Table 1 Relative length of the proximal pollical phalanx in some anthropoids Length of proximal phalanx I/length of proximal phalanx II*100 Human Gelada Baboona Gorilla Bonobo Chimpanzee Orangutan
a b

109

n 19 7 5 10 10 17 10

s.d. 1.61 1.97 0.89 4.67 2.18 3.66 2.20

76 73b 65 55 55 53 39

The baboon in this sample is P. ursinus. The high value for thumb/index ratio in geladas results from a short index nger rather than a long thumb.

primates, a biconcave basal articular surface for articulation with the bipartite distal trochlea of the proximal phalanx is apparent (Fig. 5). Also characteristic of this basal articular morphology on the middle phalanges, but not evident in Fig. 3b of Moya ` -Sola ` et al., is a beak-like process centrally located on the palmar rim of the basal articular collar (Fig. 5). Features typical of middle phalanges can be seen on the purported pollical proximal phalanx of IGF 11778 (Moya ` -Sola ` et al., Fig. 3b; Fig. 5 here). Also on the fossil gured by Moya ` -Sola ` et al. as a pollical proximal phalanx of IGF 11778 are insertion sites for the bifurcate tendon of the m. exor digitorum supercialis (FDS) and the brous exor sheath (Fig. 5). These features are characteristics of middle phalanges, not pollical proximal phalanges, as the FDS does not insert on the pollical proximal phalanx. A true pollical proximal phalanx possesses a characteristic asymmetrical, bilateral marking on the volar surface for the attachment of the oblique pulley (Doyle and Blythe, 1977). Finally, the reader will note that the hand of IGF 11778 displayed in Fig. 3b (Fig. 4b here) is missing one middle phalanx. In this gure the misplaced middle phalanx (most likely III based on its length compared to the others) has been positioned in the space where the pollical proximal phalanx should be [for a reconstruction of IGF 11778 dierent from that of Moya ` -Sola ` et al. and like that oered here, see Begun (2002)]. As a result of mistaking a nger middle phalanx for a thumb proximal phalanx, Moya ` -Sola ` et al.

calculated erroneously long thumb lengths for Oreopithecus. In addition, the incompleteness of the distal phalanges makes it dicult to discern in BA #140 whether or not the distal phalanx identied as pollical is, in fact, that of a thumb. Across primate taxa the most reliable evidence of whether a distal phalanx is that of a thumb or that of a nger is found in the shape of the base. The bases are incomplete in two of the three BA #140 distal phalanges. With respect to their overall shape, neither of the purported pollical distal phalanges (phalanx BA#130 or that of IGF 11778) bears a morphological resemblance to humans or chimpanzees, as Fig. 4 of Moya ` -Sola ` et al. demonstrates. Susman (1988) has shown that relative size and shape of the apical tuft of the distal phalanx distinguishes humans from apes, with humans having large apical tufts. The tufts of neither BA #130 nor IGF 11778 reveal humanlike proportions. Nor do the distal phalanges of BA #130 and IGF 11778 (Fig. 2), with their narrow tufts and bodies, recall morphology seen in hominids (including Australopithecus spp., Paranthropus, Homo habilis, and H. sapiens). There are two possible explanations for the ndings in Moya ` -Sola ` et al. (1999). The rst is that Oreopithecus and some papionins in Moya ` -Sola ` et al.s sample have thumb and index proximal phalanges of equal length and Oreopithecus has the longest relative thumb of any fossil or living primate, including humans. The second explanation is that the two phalanges on which Moya ` -Sola ` et al. based their regressions are not those of

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Fig. 4. Hands of Oreopithecus (from Moya ` -Sola ` et al., 1999, Fig. 2a and b). BA #140 is the partial right hand of a sub-adult individual. IGF 11778 is the partial left hand of an adult. The bones circled are not, in fact, proximal phalanges of the thumbs (see text for explanation). Metacarpal III (left) is upside down.

thumb and index nger. Both the metric and morphological data presented by Moya ` -Sola ` et al. suggest that the latter is the case.

Hand length of Oreopithecus An essential characteristic of the locomotor and postural mechanism of primates is hand length. Moya ` -Sola ` et al. (1999: 313).

Moya ` -Sola ` et al. propose that a short hand length relative to both body weight and humerus length characterizes IGF 11778 and interpret this as further evidence of precision grasping in Oreopithecus. Fig. 1 in Moya ` -Sola ` et al. (1999) employs a body weight of 32 kg for IGF 11778 (Jungers, 1987, 1990). When Moya ` -Sola ` et al. regress hand length on body weight, Oreopithecus has a short hand, with a length comparable to that of humans, gorillas, and Proconsul. Most apes, with relatively

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Fig. 5. Middle phalanx III of (a) human, (b) chimpanzee, and (c) Oreopithecus (IGF 11778). Proximal phalanx I of (d) human and (e) chimpanzee. Oreopithecus is taken from Moya ` -Sola ` et al. (1999). Arrows indicate the attachments of the exor sheaths (A4 pulleys) in (a), (b), and (c). Arrows in (d) indicate the attachment of the oblique pulley. Note also the impressions for the exor digitorum supercialis insertions medial to the exor sheath ridges on (a), (b), and (c). The middle phalanges (a, b, and c) have bipartite basal articular surfaces. Proximal phalanges, including (d) and (e), have a single basal articular surface.

longer hands, cluster on an elevated regression line. To estimate hand length, Moya ` -Sola ` et al. add together the lengths of the carpals, metacarpals, and phalanges of the third digit. From the position of Oreopithecus on the Y-axis in their Fig. 1a, IGF 11778 has a hand length of approximately 178180 mm. While it is unclear which of the IGF 11778 metacarpals and phalanges the authors include in the third ray, my measurements of the lengths of the lunate (ca. 15 mm), the deformed capitate (ca. 14 mm), third metacarpal (65.0 mm), third proximal phalanx (47.6 mm), third middle phalanx (34.7 mm), and third distal phalanx (21.0 mm) creates a hand length of over 195 mm, which positions IGF 11778 at the elevation of the ape line of Moya ` -Sola ` et al. (their Fig. 1a). In neither case do Moya ` -Sola ` et al. provide the condence limits of their regressions. As for relative hand length, hand length index in Oreopithecus is 32 (hand length/humerus+radius length 100) (Susman, 1985), exceeding averages for all great apes including gorillas (Schultz, 1960),

chimpanzees, and even orangutans (Jungers, 1985)2 [data from Napier and Napier (1967)]. Two caveats bear repeating with regard to hand length. First, and most obvious, is the problematic attempt to calculate hand length based on fragmentary, distorted fossils. Second, and most important, is the potential for error introduced by misassignment of individual hand elements.

What does the hand of Oreopithecus tell us about this Miocene ape? The hand of Oreopithecus has been studied by a number of investigators over the years (Hu rzeler, 1958; Straus, 1963; Szalay and Delson, 1979; Susman, 1985; Harrison, 1986, 1991; Begun, 2002).
2 The value of 28 for the relative hand length of orangutans, less than expected for the most arboreal of the great apes, reects the relatively long humerus and radius in this species (Jungers, 1987).

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Fig. 6. IGF 11778 metacarpals in prole. The heads of IIIV are mediolaterally attened and their articular surfaces eroded. Metacarpal I and the proximal portion of V were not recovered. Bar=1 cm.

Earlier discussions of the functional morphology of the hand in Oreopithecus have noted the poor condition of the fossils (Straus, 1963) and thus inferences have tended to be conservative. As for the fossils, none of the authors who have studied the Oreopithecus remains, including BA #140 and IGF 11778, have identied a complete set of thumb bones. Neither Susman (1985) nor Begun (2002) identied pollical metacarpals or pollical proximal phalanges in the IGF 11778 hand. Harrison (1986: 554) noted that the metacarpals of IGF 11778 were mediolaterally crushed (Fig. 6). As for elements of other thumbs, Harrison (1991: 239) states The thumb appears to have been quite short and stout, but highly mobile. As for the ngers he adds, The lateral manual digits are long, quite slender and ventrally curved. Harrisons account of both the thumb and ngers

distinguishes Oreopithecus from humans and likens it to extant apes. Harrison based his remarks on 15 metacarpals and 117 isolated phalanges. These included 2 pollical phalanges. He did not report nding a pollical proximal phalanx among the IGF 11778 hand bones. Despite the generally poor state of preservation of the individual bones, some morphology of the wrist and ngers can be interpreted and compared with anthropoid primate counterparts. As others have found, many discernible features support the notion of a large-bodied arboreal ape emphasizing power grasping rather than precision grasping in the hand. Szalay and Delson (1979: 434) described the hand of IGF 11778 as elongate and gracile and concluded that . The long hand and mobile elbow joint indicated climbing ability. Szalay and Delson thought that Oreopithecus made extensive

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Fig. 7. Proximal phalanx V of IGF 11778. Pph V is the best preserved of the proximals and yields the most reliable included angle (longitudinal curvature), which is 36(. Phalangeal curvature places Oreopithecus within the African ape range, and outside the range of humans. Great apes, including the quintessentially arboreal orangutan, have curved ngers for power grasping. Humans have short, straight ngers in keeping with their precision grasping proclivities. Vertical lines are group means, black bars are 95% condence limits of the means, and long bars are 95% condence limits of the population. For a discussion of phalangeal curvature, see Susman et al. (1984) and Stern et al. (1995).

use of the forelimb for arm-swinging and climbing. Harrison (1986: 554), in the most exhaustive study of Oreopithecus, described the metacarpus of IGF 11778 as exhibiting . a degree of curvature typical of arboreal primates and . manual phalanges that were long and recurved, suited for an arboreal habitat. My assessment of phalangeal curvature in IGF 11778 accords with Harrisons. Curvature of proximal phalanx V of IGF 11778, the best preserved of the four, is like that of African apes (Fig. 7).3 There are at least two well-preserved scaphoid bones of Oreopithecus. The scaphoid is particularly relevant to considerations of thumb proportions because the scaphoid serves as the base of support
Three measurements are used to derive the included angle (=longitudinal curvature) of the proximal phalanx. They are 1) interarticular length, 2) dorsopalmar midshaft diameter, and 3) distance of the dorsalmost point at midshaft above the interarticular baseline (see Fig. 1, Stern et al., 1995). Included angle is calculated with these three measurements (see Susman et al., 1984; Stern et al., 1995).
3

for the trapezium and the long bones of the thumb itself. Hominoids with the longest thumbs, in order, humans, gorillas, bonobos, and chimpanzees, have correspondingly robust scaphoid bones with fused os centrales. In contrast, the scaphoid of Oreopithecus is not robust but instead is gracile, like that of orangutans. The left scaphoid of IGF 11778 is intact (Fig. 8a). Both IGF 11778 and another, smaller scaphoid (BA #26), reveal unfused os centrales as in orangutans, hylobatids, and monkeys. It is apparent, as noted by Harrison (1986), that both Oreopithecus scaphoids (IGF 11778 and BA #26) resemble their counterparts in orangutans (Fig. 8b). In humans, the os centrale normally fuses with the scaphoid early in lifeanywhere from the prenatal into the juvenile period (Fig. 8d). In African apes, the scaphoid and os centrale normally fuse later in life (Fig. 8c). In Asian apes and typically in monkeys, the os centrale remains a separate bonedistinct from the scaphoid. An unfused os centrale represents the primitive condition, common to all but a few

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Fig. 8. Scaphoid bones of (a) Oreopithecus (IGF 11778), (b) orangutan, (c) chimpanzee, and (d) human (proximal view). Oreopithecus and orangutans have an unfused os centrale. The scaphoids of humans and chimpanzees (and other African apes) have a fused os centrale (indicated by the arrow) which expands the articular surface for the trapezium (above the arrow) and trapezoid. The trapezium, in turn, is where the thumb articulates with the wrist. Humans and chimpanzees have expanded scaphoids and concomitantly larger and longer thumbs than orangutans and, presumably, Oreopithecus.

primates. As a result of the fusion of the os centrale, humans have a distinctive scaphoid that is quite dierent from that of Oreopithecus and most other primates. While many authors have suggested that fusion of the os centrale in humans may relate to increased mid-carpal stability during wrist extension, it is equally likely that the fused os centrale and its union with the scaphoid are related to the enlarged thumb in humans. The fusion of the os centrale serves to strengthen the scaphoid neck and tubercle, and the fusion enhances the surface that supports the trapezium. The trapezium, in turn, supports the robust pollical metacarpal of the human thumb. The lack of a fused os centrale in Oreopithecus runs counter to the idea of a hominidlike conguration of the base of the thumb. Orangutans, which also have an unfused os centrale and weakly developed scaphoid neck and tubercle, concomitantly have the shortest thumbs and are perhaps the least capable precision graspers among the hominoids. Moya ` -Sola ` et al. (1999) infer that Oreopithecus had a strong exor pollicis longus muscle (FPL). In their Fig. 4b, which depicts the palmar view of BA #130, much of the bone that would normally serve as the insertion site for the tendon of the FPL is missing. Next to it, in their Fig. 4c, lies the pollical distal phalanx of IGF 11778. Neither Harrison (1986) nor I have noted the presence of an FPL insertion on the pollical distal phalanx of

IGF 11778. The question of a FPL in Oreopithecus is, at best, indeterminate. Finally, while Straus (1949, 1963) and Susman (1994, 1998) have noted that humans alone among primates have a true FPL, there are others who argue that the presence of an FPL is not diagnostic of humanlike precision grasping (Marzke, 1997).

Other features of Oreopithecus that bear on the question of thumb and hand length The limb proportions of Oreopithecus have important implications for reconstructing hand length and proportions of the thumb and ngers because there are no higher primates that have long arms and forearms with short hands and ngers. Hu rzeler (1958) concluded that Oreopithecus sported long forelimbs and short hindlimbs, resulting in a high intermembral index (see Fig. 1). Schultz (1960), Straus (1963), Susman (1985), Harrison (1986), Jungers (1987, 1990) and Rose (1993) all have commented on the high intermembral index, i.e., long forelimb and short hindlimb, of IGF 11778. Among hominoids a high intermembral index represents the interlimb conguration of large-bodied climbers (Cartmill, 1974; Jungers, 1985). Schultz (1960) thought that Oreopithecus was an arboreal climber similar to gorillas. Straus (1963) concluded from limb

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proportions that Oreopithecus had a high intermembral index indicative of a brachiator. Schultz (1960), Straus (1963) and Harrison (1986) also commented on articular dimensions of IGF 11778 and noted it had a large humeral head diameter, exceeding that of the femur. The humerofemoral head diameter index reects the high humerofemoral length index in Oreopithecus and also falls within the ape range. Jungers (1987, 1990) studied body size and skeletal proportions of Oreopithecus and calculated a body mass of 32 kg for IGF 11778. Jungers (1987) concluded that the only great ape with a relative forearm length as great as that estimated for Oreopithecus was the orangutan. Using a principle coordinates analysis of average taxonomic distances based on mass-adjusted variables of the limbs, Jungers concluded that the limb proportions of Oreopithecus were most similar to those of female bonobos and female orangutans. When a narrow allometric comparison was conducted with primates of similar body size, Oreopithecus exhibited a longer humerus and a longer radius than chimpanzees or bonobos. The upper limb of Oreopithecus was also considerably longer than that of like-sized baboons. A clustering routine revealed that when humerus length, radius length, femur length, tibia length, and ilium length were taken into account, Oreopithecus clustered with orangutans (Jungers, 1990). As did Biegert and Maurer (1972), who studied limb proportions earlier and likened Oreopithecus to orangutans, Jungers concluded that body proportions of Oreopithecus indicated a predominance of climbing and brachiation rather than pronograde quadrupedalism.4

Susman (1985) also concluded, based on a study of selected, better-preserved parts of the IGF 11778 skeleton, that Oreopithecus was arboreal and apelike in its postcranium. Combining 1) the high hand length index, 2) a qualied determination of phalangeal curvature, and 3) assessment of interlimb proportions, Susman suggested that Oreopithecus was as well-adapted to suspensory positional behavior as extant chimpanzees, but perhaps less well-adapted for arboreality than extant orangutans. All evidence of limb proportions supports the notion that Oreopithecus was an arboreal hominoid with interlimb proportions that would have made it an eective, large-bodied, climbing, suspensory-adapted animal, not unlike apes of today. Consistent with the above are a long hand that possessed long, curved ngers and an orangutan-like scaphoid. Given a consensus of almost all researchers on the high intermembral index and apelike positional behavior of Oreopithecus, it would be entirely inconsistent to nd a hand with humanlike proportions. Such models, viz. anthropoids with long arms, long forearms, but short hands, do not exist today, and there is no evidence that they ever existed in the past. There is no convincing evidence that Oreopithecus possessed a short hand with a long, humanlike, precision grasping thumb. There is no evidence that supports the notion of humanlike precision grasping in Oreopithecus. Rather, the morphological design of the hand is consistent with the idea that Oreopithecus lived in trees and as such, like most primates, and especially the largebodied hominoids, relied on power grasping more than precision grasping.

4 With a high intermembral index that includes a short hindlimb (Hu rzeler, 1960; Schultz, 1960; Straus, 1963), 6% shorter than that of African apes (Jungers, 1987), a long ilium, a foreshortened midfoot, an abducted hallux, a long hand with curved ngers, the possibility that Oreopithecus was a biped (Ko hler and Moya ` -Sola ` , 1997; Rook et al., 1999) is as remote as the notion that Oreopithecus had a humanlike precision grasp. Nor does the conclusion based on faunal and oral considerations that Oreopithecus inhabited a swampy forest (Azzaroli et al., 1986) with tree stands at some distance (Weyland, 1962) support the idea that Oreopithecus was a biped.

Acknowledgements Thanks to Annalisa Berzi, Augusto Azzaroli, and Giovanni Ficcarelli for permission to study original Oreopithecus fossils in Florence. Thanks to Dr. Eric Delson and Dr. Terry Harrison for providing access to casts of IGF 11778 and measurements of additional Oreopithecus fossils. Comparative primate material was measured at the American Museum of Natural History. Data

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R.L. Susman / Journal of Human Evolution 46 (2004) 105117 Jungers, W.L., 1990. Problems and methods in reconstructing body size in fossil primates. In: Damuth, J., MacFadden, B.J. (Eds.), Body Size in Mammalian Paleobiology: Estimation and Biological Implications. Cambridge University Press, Cambridge, pp. 103115. Ko hler, M., Moya ` -Sola ` , S., 1997. Ape-like or hominid-like? The positional behavior of Oreopithecus bambolii reconsidered. Proc. Natl. Acad. Sci. 94, 1174711750. Landsmeer, J.M.F., 1962. Power grip and precision handling. Ann. Rheum. Dis. 21, 164170. Marzke, M.W., 1997. Precision grips, hand morphology, and tools. Am. J. Phys. Anthrop. 102, 91110. Moya ` -Sola ` , S., Ko hler, M., Rook, L., 1999. Evidence of hominid-like precision grip capability in the hand of the Miocene ape, Oreopithecus. Proc. Natl. Acad. Sci. 96, 313317. Napier, J.R., 1956. The prehensile movements of the human hand. J. Bone Jt. Surg. Am. 38, 902913. Napier, J.R., Napier, P.H., 1967. A Handbook of the Living Primates. Academic Press, London. Rook, L., Bondioli, L., Ko hler, M., Moya ` -Sola ` , S., Macchiarelli, R., 1999. Oreopithecus was a bipedal ape after all: evidence from the iliac cancellous architecture. Proc. Natl. Acad. Sci. 96, 87958799. Rook, L., Renne, P., Benvenuti, M., Papini, M., 2000. Geochronology of Oreopithecus-bearing succession at Baccinello (Italy) and the extinction pattern of European Miocene hominoids. J. Hum. Evol. 39, 577582. Rose, M.D., 1993. Functional anatomy of the elbow and forearm in primates. In: Gebo, D.L. (Ed.), Postcranial Adaptation in Nonhuman Primates. Northern Illinois University Press, DeKalb, pp. 252272. Sarmiento, E.E., 1987. The phylogenetic position of Oreopithecus and its signicance in the origin of the Hominoidea, Am. Mus. Novitates 2881, New York, pp. 144. Sarmiento, E.E., Marcus, L.F., 2000. The os navicular of humans, great apes, OH 8, Hadar, and Oreopithecus: function, phylogeny, and multivariate analyses. Am. Mus. Novitates 3288, New York, pp. 138. Schultz, A.H., 1960. Einige beobachtungen und masse am skelett von Oreopithecus im vergleich mit anderen catarrhinen primaten. Z. Morph. Anthrop. 50, 136149. Straus, W.L., 1949. The riddle of mans ancestry. Qtr. Rev. Biol. 24, 200223. Straus, W.L., 1963. The classication of Oreopithecus. In: Washburn, S.L. (Ed.), Classication and Human Evolution. Aldine, Chicago, pp. 146177. Stern, J.T., Jungers, W.L., Susman, R.L., 1995. Quantifying phalangeal curvature: an empirical comparison of alternative methods. Am. J. Phys. Anthrop. 97, 110. Susman, R.L., 1985. Functional morphology of the Oreopithecus hand. Am. J. Phys. Anthrop. 66, 235. Susman, R.L., 1988. Hand of Paranthropus robustus from Member 1, Swartkrans: fossil evidence for tool behavior. Science 239, 781784. Susman, R.L., 1994. Fossil evidence for early hominid tool use. Science 265, 15701573.

on human hands come from the Cleveland Museum of Natural History. Thanks to Lucille Betti-Nash for preparing the gures. John Fleagle, Bill Jungers, Jack Stern, Theresa Franz, and Biren Patel provided comments on the manuscript.

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