High occurrence of a basicranial feature in Homo erectusAnatomical description of the preglenoid tubercle.код для вставкиСкачать
THE ANATOMICAL RECORD (PART B: NEW ANAT.) 274B:148 –156, 2003 FEATURE ARTICLE High Occurrence of a Basicranial Feature in Homo erectus: Anatomical Description of the Preglenoid Tubercle VALERY ZEITOUN* The degree of differentiation between Western and Eastern Early Middle Pleistocene hominids (Homo erectus) is still under debate. On the one hand, the two populations are believed by some to belong to two separate taxonomic categories possessing their own exclusive features. On the other hand, they are considered by others to belong to a single but ubiquitous global population with regional specificities that are mainly due to variations in the frequency of certain features. In this context, an additional trait, the preglenoid tubercle (tuberculum anterius fossae mandibularis), may well shed light on the matter. The tubercle is often present on the anterior part of the fossa mandibularis at the anterior extent of the joint capsule and articular tubercle (tuber articulare) of fossils usually assigned to Homo erectus. This study describes the anatomy of the preglenoid tubercle, to indicate its occurrence and to try to improve its use in taxonomy and phylogeny, although its origin and functional significance have not yet been clarified. Anat Rec (Part B: New Anat) 274B:148 –156, 2003. © 2003 Wiley-Liss, Inc. KEY WORDS: Homo erectus; temporomandibular joint; preglenoid tubercle; fossil; bone INTRODUCTION There is an ongoing debate in paleoanthropology concerning whether the Early and Middle Pleistocene Homo erectus should include specimens from across Eurasia and Africa or should be limited to specimens from eastern Eurasian (Rightmire, 1984; Dr. Zeitoun has a master degree in geology. He obtained his PhD in Physical Anthropology in 1996. As a paleoanthropologist his research interest is the reappraisal of the definition of the species Homo erectus using cladistics. Since 1992, he has been working in Indonesia in collaboration with T. Jacob and since 1999, he collaborated on a three-dimensional imaging program with J.J. Hublin. An Assistant Professor for 2 years in the Chaire de Préhistoire et Paléoanthropologie in College de France, and since 2001, he is a Researcher in CNRS at the UPR 2147 in Paris. He is the director of the Thai–French Paleosurvey, looking for Pleistocene human remains in the caves of Northern Thailand. *Correspondence to: Valery Zeitoun, 44 rue de l’Amiral Mouchez, 75014 Paris, France. Fax: ⴙ33-143-1356-30; E-mail: firstname.lastname@example.org DOI 10.1002/ar.b.10028 Published online in Wiley InterScience (www.interscience.wiley.com). © 2003 Wiley-Liss, Inc. Hublin, 1986; Turner and Chamberlain, 1989; Brauer and Mbua, 1992; Kennedy, 1991). If the latter approach is taken, the western Old World members of Early and Middle Pleistocene hominids would be assigned to “Homo ergaster ” (Andrews, 1984; Groves, 1989; Wood, 1992; Stringer, 1996). The “two-species” interpretation maintains that the eastern Eurasian members of this taxonomic group (see Box 1 ) exhibit a suite of derived characters relating to ancestral African members of the genus Homo. The traditional one-species interpretation argues principally that any differences between the remains attributed to eastern and western populations are due to a combination of normal, within-species variations. These variations in turn are due to a combination of temporal spread and isolation by distance. Many of the traits used to separate Early and Middle Pleistocene hominid eastern and western populations appear to differ between regions more in terms of frequency and degree of expression than in terms of presence or absence. Independently of the strictly taxonomic issues, the degree of differentiation between western (principally African) and eastern (principally Southeast Asia) Old World sequences of Early and Middle Pleistocene archaic Homo raises questions regarding the degree and pattern of hominid differentiation after the first major geographical dispersal of hominids in the Early Pleistocene. Additional paleontological data that might have a bearing on these issues are therefore of interest. In this context, an additional morphological feature, the preglenoid tubercle (tuberculum anterius fossae mandibularis) is described and discussed with respect to its distribution through hominid evolution. SPECIMENS EXAMINED Initially, presence/absence of a tubercle was recorded on the anterior part of the temporomandibular joint within the original fossil human specimens from the Ngawi, Ngandong, Sangiran, and Sambungmachan sites in Indonesia (Figures 1– 4). For the basis of further comparisons, observations on the tubercle of Australopithecines and several Homo speci- FEATURE ARTICLE THE ANATOMICAL RECORD (PART B: NEW ANAT.) 149 Box 1: Glossary of Terms Taxon (singular, Taxa plural): A group of organisms that are recognized as a formal unit at each level of the classification (Simpson, 1961). A group of organisms that is given a name (Wiley et al., 1991). Paraphyletic groups: groups for which one or more descendants of an ancestor are excluded from the group (Wiley et al., 1991). Hyperostotic traits: traits for which there appears an excess of bone growth (Hauser and De Stephano, 1989). Apomorphic character: it is the descendant character by opposition to the ancestral character a feature is apomorphic (Wiley et al., 1991). Autapomorphic: a feature is autapomorphic if it is apomorphic and if it is not shared by other taxa than a single taxon (Darlu and Tassy, 1993). mens from across the Old World (Europe, Asia, and Africa) were made on original specimens and high-quality casts of them. Designations of fossil Homo specimens in this study are given (Table 1) for the convenience of common referral and do not reflect my own taxonomic assignments. Presence/absence of this tubercle was also recorded on the skull of male and female Pan, Gorilla, and Pongo (Collection of Museum National d’Histoire Naturelle, Paris and National Museum of Natural History, Leiden; Table 2) and in 1,500 adult modern human individuals sampled globally (collection of Musée de l’Homme, Paris; Table 3). ANATOMICAL DESCRIPTION OF THE PREGLENOID TUBERCLE In norma basilaris (Figure 5), a tubercle can be seen on the anterior part of the fossa mandibularis, at the anterior limit of the glenoidal fossa of the temporomandibular joint, at the anterior extent of the joint capsule on the tuber articulare. The tubercle has a rounded conic shape with a diameter of approximately 1.1 mm at the bottom with a height of approximately 0.3 mm. It is proposed here that this tubercle be called the tuberculum anterius fossae mandibularis or preglenoid tubercle. According to what is shown in anatomical literature, it should be the insertion point of the short tendinous bunches that is linked to the superior head of the musculus pterygoideus lateralis before the anterior edge of the discus articularis. Unfortunately, these short tendinous bunches are not described by all authors who make the description or study of this anatomical area (Paturet, 1951; Rouvière, 1954; Burch, 1970; Griffin et al., 1975; Schmolke, 1994; Abe et al., 1997). Nevertheless these bunches are illustrated and called anterior temporomeniscal fibres of the musculus pterygoideus lateralis (Figure 6) by Nicolas et al. (1926). OCCURRENCE OF THE PREGLENOID TUBERCLE The preglenoid tubercle is present in the main Homo erectus fossils from Indonesia. It is present in all fossils of the Ngandong series whenever the anatomical area is preserved. It may be observed in the Sangiran 2, 10, 17, Ngawi 1, and Sambungmachan 1 specimens (the anatomical area is absent in Sambungmachan 3). The size of the tubercle is constant for all the Asian specimens. It is noticeable that the preglenoid tubercle is not present on the temporal bone of Sangiran 26. In Africa, among specimens assigned to or closely affiliated with Homo erectus, the preglenoid tubercle is only present in KNMER 3883. This is one of the earliest African Homo erectus specimens (Rightmire, 1990; Wood, 1991). The tubercle here is more pronounced than on any Indonesian specimen (approximately five times larger in volume). Indeed, at the bottom, the diameter is a little bit less than two times the diameter of Indonesian specimens when measured on enlarged picture. The casts of the juvenile specimen KNM-WT 15000 seem to show a little knob at the position of the preglenoid tubercle, but, as a result of taphonomical disturbance, the preservation of the temporal bone is not good enough to con- firm this (Prat, in a personal communication, has the opportunity to check the original specimen and assesses that the feature is absent). The preglenoid tubercle is absent in Great Apes and very rare in modern humans. However, the reservation has to be made that there is no possible confusion between the large size of the tuberculum anterius fossae mandibularis in Homo erectus and what has been observed in 2.7°/°° of modern humans (Table 3), where the volume is evaluated to be approximately 20% of what may be seen in Homo erectus. Rather than a tubercle, it is an extremely slight vestigial knob at the insertion point of the anterior temporomeniscal fibres up to the superior head of the m. pterygoideus lateralis (Figure 6). Unfortunately, the anatomy of the temporomandibular joint capsule and its possible relationships to other structures near the joint are not fully understood. HYPOTHESES SURROUNDING THE TUBERCLE If the simplest anatomical hypothesis is that the tuberculum anterius fossae mandibularis is the bony insertion point of the anterior temporomeniscal fibers up to the superior head of the musculus pterygoideus lateralis, the next questions to ask are what is the function of these fibers and how do they work. Unfortunately, the anatomy of the temporomandibular joint capsule and its possible relationships to other structures near the joint are not fully understood (Schmolke, 1994). Opinions vary in the literature as to the role of the ligaments associated with the temporomandibular joint. In a bibliographic survey of temporomandibular joint function, Sato et al. (1995) noted that the functional role of the temporomandibular joint ligaments is 150 THE ANATOMICAL RECORD (PART B: NEW ANAT.) Figure 1. Right temporal bone of Ngandong 12 in norma basilaris. The anterior part of the skull is to the right. poorly known and that additional investigation is needed. The same applies to the exact role of the lateral pterygoid muscle, especially in controlling horizontal jaw movements (Murray et al., 2001). Although it is well known that (1) the joint capsule has primarily an anterior relationship with the lateral pterygoid muscle and (2) the latter muscle has two heads that exhibit functionally reciprocal activation, the exact insertion point and function of the superior head of the lateral pterygoid muscle remains controversial (Piette, 1993; Murray et al., 2001). The existence and the functional role of inconsistent associated structures such as the anterior temporomeniscal fibres are difficult to assess. However, according to Griffin et al. (1975), the ligament of the lateral pterygoid muscle appears to be the restraining ligament for the joint. Its fibres both draw the articular disc from the anterior side and balance it by supporting it from the posterior side. The ligament fibres attached to the articular disc draw the disc from the posterior side in its course of mandibular closing, thus enabling the articular disc to move smoothly (Abe et al., 1997). At the same time, the shaping of the articular cavities and the texture of the joint capsule allow movements of the articular disc, predominantly in the anteromedial direction (Schmolke, 1994). This latter author specifies that, on the entire medial side of the joint, the articular disc and its capsular attachments are in close contact with the fascia of the musculus pterygoideus lateralis whereby a small portion of the superior head of this muscle inserts directly into the anteromedial part of the discus articularis. It is not said that there is any insertion on the tuberculum articulare that could confirm the nature of the preglenoid tubercle and there is no mention of differences concerning the occurrence or the strength of the anterior temporomeniscal fibres of the musculus pterygoideus lateralis as illustrated in Nicolas et al. (1926). The same is true of the compilation of epigenetic variants of the human skull published by Hauser and De Stefano (1989), where nothing is said on this subject. Because both the superior and the inferior heads of the musculus pterygoideus lateralis are agreed upon to influence the position of the articular disc during temporomandibular joint movements (Schmolke, 1994), the proximity of the location of the anterior temporomeniscal fibres should give some clue on its role. It is also reported that, apart from viscoelastic deformation, the other important disc stabilisation mechanism seems to be related to attachments of the disc (Piette, 1993). Moreover, particular structural properties of disc attachment are due to the morphology of the human temporomandibular joint ligament that is ligamentous without distinct structure and the sheath-like FEATURE ARTICLE structure and other components of the lateral ligaments store energy and protect the capsule from stress and tension during movements of the jaw (Sato et al., 1996) and could transfer power until the fibres insertion. Thus, according to Sato et al. (1996) and Schmolke (1994), the preglenoid tubercle may be an invariant superior attachment of the anterior temporomeniscal fibres on the anterior part of the discus articularis that can play a role in the anteromedial mechanism of that area of the temporomandibular joint. As a general and common property, it can be proposed that the strength of the preglenoid tubercle is at least indirectly linked to the activity of the short tendinous fibres of musculus pterygoideus lateralis. Three hypotheses may be considered to explain the occurrence of the preglenoid tubercle: (1) hyperostotic as a genetic inheritability linked to the global robustness of Homo erectus as formerly proposed by Hublin (1989); (2) hyperostotic as a biomechanical function due to a specific geometry of the temporomandibular joint among Homo erectus; (3) hyperostotic as a pathological excrescence. Genetic Inheritability and Global Robustness If the preglenoid tubercle is a developed insertion point among others, some hyperostotic changes should be observed in the neighbouring cranial area. This is the case in the Ngandong series, where the base of the skull dis- Figure 2. Left temporal bone of Sambungmachan 1 in norma basilaris. The anterior part of the skull is to the right. FEATURE ARTICLE THE ANATOMICAL RECORD (PART B: NEW ANAT.) 151 Figure 3. Left temporal bone of Ngawi 1 in norma basilaris. The anterior part of the skull is down to the right. or lesser obliquity of the posterior slope of the tuberculum articulare; the proportional length or breadth of the anterior wall of the fossa mandibularis; the length of a subtemporal plane. This list of features is linked with biomechanical properties of the temporomandibular articulation. Thus, it seems that, in biomechanical terms, there is no morphological particularity in Homo erectus that might be linked to the occurrence of the preglenoid tubercle due to specific biomechanical properties. If there is no structural pattern that suggests the presence of the tubercle, the distinction between presence and absence of preglenoid tubercle should be at another level, as a geographic specificity, may be in terms of inheritability on soft tissues. That point could support the interpretation of Brauer and Mbua (1992) regarding Homo erectus vs. Homo ergaster as two different populations, but it is very difficult to prove. It can also be due to a particular activity that may intimately be tied to diets. The answer to that hypothesis should be found in the environment as far as it can allow us to distinguish Indonesian, Chinese, and most of the African specimens. Biomechanical Function A review of different parameters relevant to the anatomy of the temporomandibular articulation found by Picq (1983) and Zeitoun (1996) among all the specimens described Table 1 does not show any correlation to the presence of a preglenoid tubercle. To sum up these parameters, there is no relationship between the occurrence of the tuberculum anterius fossae mandibularis and the following: the greater or lesser contribution of the tympanal plate to the posterior wall of the fossae mandibularis; the existence of a big or small postglenoidal process; the occurrence of a postglenoidal process transversally extended or laterally offset; the greater The tuberculum anterius fossae mandibularis could serve as a strong anterior brake for the temporomandibular joint (Picq, personal communication). It is noteworthy, that a lengthening contraction of the musculus pterigoideus lateralis has the effect of slowly letting out a rope controlling the condyle as it travels back into the fossa (Wilkinson, 1988). Because the flatness of the anterior portion of the temporomandibular joint fails to provide a bony buttress against movements, a stronger muscular insertion has to be present for compensation. In this view, the flatness of the temporomandibular joint could be seen as a lack of anatomical protection for the anterior part of the tympanal plate and would need to be compensated by strong fibres. Figure 4. Left temporal bone of Sangiran 2 in norma basilaris. The anterior part of the skull is to the right. plays hyperostotic clues such as a well-developed precondylar tubercle, a pharyngeal tubercle, a peribasic basal excrescence or postcondyloid tuberosities (Figure 5 and also Durband, 2002). After Hublin (1989), all Homo erectus id est, including African and Chinese have a global robusticity of their skeleton nevertheless almost only the Indonesian specimens possess this particularity. In other respects, it should be stressed that almost all fossil and living human and Great Ape taxa with hyperostotic features do not exhibit any tuberculum anterius fossae mandibularis. Pathologic Excrescence Elyaqtine (in an oral communication) informs me that a weak preglenoid tubercle, in association with a patho- 152 THE ANATOMICAL RECORD (PART B: NEW ANAT.) FEATURE ARTICLE TABLE 1. Occurrence of the preglenoid tubercle among fossils Australopithecines Kenya Kenya Kenya Ethiopia Ethiopia South Africa South Africa South Africa South Africa South Africa Homo sp. Ethiopia Kenya Homo habilis Kenya Homo rudolfensis Kenya Homo erectus Indonesia Indonesia Indonesia Indonesia Indonesia Indonesia Indonesia Indonesia Indonesia Indonesia Indonesia Indonesia China China Occurrence Homo ergaster °KNMER 406 °KNMER 407 a °KNMER 732 *AL 33345 °OH 5 °Sts 5 °Sts 71 °Stw 505 °SK 47 °SK 48 N N N N N N N N N N °OH 24 °KNMER 1805 N N °KNMER 1813 N °KNMER 1470 N °Ngandong 1 °Ngandong 6 °Ngandong 7 °Ngandong 10 °Ngandong 12 °Ngawi 1 °Sambungmachan 1 °Sangiran 2 °Sangiran 4 °Sangiran 10 °Sangiran 17 °Sangiran 26 *Sinathropus III *Sinathropus XI Y Y Y Y Y Y Y Y Y Y Y Y Y Y Kenya Kenya Kenya Homo sapiens Archaic Tanzania Tanzania Kenya Ethiopia Zambia Morocco India China Modern France Near East Near East Neandertal Near East Greece Italy Italy Croatia Germany Belgium Belgium France France France Uzbekistan Occurrence °KNMER 15000 *KNMER 3883 *KNMER 3733 N Y N *LH 18 *Ndutu 1 *ES 11693 *Omo Kibish 2 °Kabwe 1 °Sale *Narmada *Dali N N N N N N N N °Malaurie *Skhul V *Qafzeh 9 N N N *Amud *Petralona *Monte Circeo 1 *Saccopastore 2 *Krapina 3 *Steinheim *Spy 1 *Spy 2 *La Ferrassie 1 *La Quina H5 *La Chapelle aux Saints *Teshik Tash N N N N N N N N N N N N Y, preglenoid tubercle present; N, preglenoid tubercle absent; *, checked on cast; °, checked on original. logic stretching of the tuberculum articulare, is present on the tardenoisian Homo sapiens sapiens from Malaurie that is published by Couture et al. (1990). In this study, this stretching is interpreted, by Crubezy, as a recurrent luxation of the mandible. This clinical argument tends to confirm the hyperostotic property, because the luxation triggers high strain on the musculus pterygoideus lateralis and also indirectly on the anterior temporomeniscal fibres. Nevertheless, it is difficult to support the hypothesis that all the Indonesian specimens could have the same pathology. It should also be stressed that almost all fossil and living human and Great Ape taxa with hyperostotic features do not exhibit any tuberculum anterius fossae mandibularis. At that step of the survey, no conclusive argument has been able to explain the evolutionary process of the preglenoid tubercle. The hyperostotic hypothesis as a clue to a particular use of the temporomandibular joint of Homo erectus specimens seems to be the best, although there is no evidence for a link with any particular bone morphology as seen in the second hypothesis. It is nevertheless clear that, insofar as the anatomical area is well preserved, no common occurrence of preglenoid tubercle exists between Homo erectus and other hominids. The problem remains as to how to define a way of understanding the significance of a feature seen on a poor fossil specimen sample TABLE 2. Occurrence of the preglenoid tubercle among great apes Gorilla gorilla* Pan troglodytes* Pongo pygmeus*° Gorilla Chimpanzee n ⫽ 26 (13 females, 13 males) Occurrence Orangutan n ⫽ 23 (12 females, 11 males) Occurrence 0 0 *Galerie d’Anatomie, Museum National d’Histoire Naturelle, Paris. °National Museum of Natural History, Leiden. n ⫽ 52 (32 females, 20 males) Occurrence Total 101 0 0 FEATURE ARTICLE THE ANATOMICAL RECORD (PART B: NEW ANAT.) 153 TABLE 3. Occurrence of the preglenoid tubercle among modern humans, Homo sapiens sapiens Modern humans Individuals Asia (Japan, Indochina, Indonesia, China, Turkey)* Africa (Morocco, Algeria, West Africa, Central Africa)* Europe (France, Russia, Romania, Italy, Germany)* America natives (Argentina, Chili, Peru)* Australia natives (Aborigines)* Total n n n n n n ⫽ ⫽ ⫽ ⫽ ⫽ ⫽ 627 472 323 60 18 1500 Occurrence 1 (Japan), 1 (Turkey) 1 (West Africa) 1 (Russia) 0 0 4 *Musee de l’Homme, Museum National d’Histoire Naturelle, Paris. through a comparison with a numerous living population where the feature is absent. Even if the problem of causation arises when differences occur between populations in the frequency of discrete traits, can the differences be attributed to biological distance, or have events played a significance? A more in-depth study is needed to understand the significance of the feature, although some works have already focused on the phylogenetic interest of discrete traits (Anthony, 1946; Falk, 1986; Barriel, 1991, 1994; Barriel and Tassy, 1991; Braga, 1995; Msuya and Harrisson, 1996). To support the use of any feature to build phylogeny, one should be reminded that problems of inheritability are relevant because discrete traits are based on phenotypic expression, and the question is to determine just how far appearances reflect their genetic base. However, this point objectively concerns all features in paleoanthropology. POTENTIAL SIGNIFICANCE When dealing with fossils, one can only make analyses from the original Figure 5. Base of the skull of Ngandong 6. database at the level of individual specimens. One of the main goals of paleoanthropology is to identify relationships between these specimens, whatever they represent (populations, species, or supraspecific taxa). Because phylogeny is supposed to reflect the history of biological events that lead to the appearance of taxa (populations, species, or supraspecific taxa), taxa are suggested by phylogeny (although the hierarchy between “clusters” needs justification, and sometimes arbitrary, and a posteriori, rules are used to divide specimens into 154 THE ANATOMICAL RECORD (PART B: NEW ANAT.) FEATURE ARTICLE Figure 6. Lateral view of the anterior temporomeniscal fibres and superior head of the musculus pterygoideus superior, as illustrated in Nicolas et al. (1926), modified. groups). Unfortunately, in many articles in paleoanthropology, an a priori grouping of the fossils is considered before any phylogenetic analysis is carried out. This kind of work can only propose a pattern of relationships between taxa that are arbitrarily defined a priori on the basis of subjective criteria, such as their common occurrence in the same locality or chronological proximity. This practice is deeply rooted in the belief that species differ from higher taxa. But we know that species are in no way different from higher taxa; at best they are higher taxa in the process formation (Løvtrup, 1987), although they can clearly be paraphyletic, as pointed out by Nelson (1989), however similar their component individuals may seem. Referring a fossil individual to a particular species before any analysis of its character distribution therefore generates a serious flaw in the taxonomic conclusions of a study. A priori groupings generate additional reversions and convergences. If, by chance, this is not the case, the implication is that one already knows which state is derived and which is primitive. Thus, strictly speaking, in the Hennigian sense, the approach suggests that one already knows the phylogeny that one is trying to reconstruct, which does not make sense. The point that has to be made is nevertheless the taxonomical level on which discussion should take place if it is possible to do so. Should the occurrence of the preglenoid tubercle be considered in terms of variation, as in Brauer and Mbua (1992), which suggests that the limit of the species is known? Or, as in As a general and common property, it can be proposed that the strength of the preglenoid tubercle is at least indirectly linked to the activity of the short tendinous fibres of musculus pterygoideus lateralis. Trinkaus (1990), should we address only the distribution of the feature, because “the delineation of the units of analysis becomes largely arbitrary, determined by relatively abundant temporal geographic gaps in the fossil record and by an a priori morphological decision as to where the breaks should occur”? A correlative alterna- tive would be just to consider the feature and take it into account before carrying out a phylogenetic analysis. If we take it that the existence of autapomorphic features for Homo erectus is unproven (Stringer, 1984; Hublin, 1986; Kennedy, 1991; Brauer and Mbua, 1992), each feature has to be taken into account, with no reason to exclude it a priori. Thus, the existence of a preglenoidal tubercle, which is common to Indonesian Homo erectus and to the KNMER 3883 specimen, which was considered as one of the first African Homo erectus (Rightmire, 1990, Wood, 1991) most commonly accepted as Homo ergaster (Groves, 1989; Wood, 1994; Strait et al., 1997), suggests regional differences in the frequency of traits within Homo erectus sensu lato but no clear-cut east– west differences. At that step of the survey on the significance of the preglenoid tubercle, its absence in Pongo, Gorilla, Pan, and Australopithecine supposes that this could be a derived state in these Homo erectus sensu lato. On the one hand, with no clear autapomorphic feature for Homo erectus species, it is not possible to assert that Homo erectus is a monophyletic species (Hublin, 1986), and at least one apomorphy needs to be found to define Homo erectus. Thus, it may be that the preglenoid tubercle appeared in Homo ergaster FEATURE ARTICLE and disappeared in Homo sapiens, or at least in the most recent Homo erectus. If African Homo erectus is taxonomically different from Asian Homo erectus as claimed by Andrews (1984), Stringer (1984), Wood (1984), and Hublin (1986), the disappearance of the preglenoid tubercle could have become a new feature (by reversion) in archaic Homo sapiens and Neandertal. On the other hand, many features can be useful in distinguishing eastern and western Homo erectus by region (Brauer and Mbua, 1992), and the preglenoid tubercle is one of these. In these terms, the trait expresses both Homo erectus and Homo ergaster as regional populations of Homo erectus sensu lato. Because of the lack of knowledge on its origin, the preglenoid tubercle should more certainly be considered as a clue to differences between populations, perhaps resulting from particular masticatory practises or dietary habits linked to the regional environment. But, as previously reported by Trinkaus (1990) and also Brauer and Mbua (1992, p. 82), who consider the specimens from Ngandong as archaic Homo sapiens (see also Dubois, 1940; Campbell, 1963; Jelinek, 1981; Tobias, 1985; Stringer, 1987; Bonde, 1989), the question of taxonomic level (is Homo erectus one single species, or shall we have to consider several geographically different species) remains unanswered. The same question arises for all the Chinese specimens, including the Sinanthropus, where the preglenoid tubercle is absent (as far as can be assessed from the remaining material— only casts made of plaster still existing). If it was certainly the case, the absence of the preglenoid tubercle could be a clue to split Chinese Homo erectus population from Indonesian Homo erectus population and to assign them to an other species. From this point of view, the problem is similar between Continental and Insular Asia and between Africa and Asia. CONCLUSION Undoubtedly further research has to be done to determine where the preglenoid tubercle is present and also to better understand its functional significance in terms of its anat- THE ANATOMICAL RECORD (PART B: NEW ANAT.) 155 omy. This last point is still unclear according to the available anatomical literature on the temporomandibular joint. At that step of the art, and going in quest of new features to shed light on phylogeny of human fossil, the occurrence of the preglenoid tubercle can be used to raise questions regarding the degree and pattern of Homo erectus. Is the tubercle more likely to be a feature that allows distinctions to be made among Homo erectus populations? Or is it a preliminary clue leading to questions as to the actual grouping of specimens at species level? At that step, it seems possible to say that this feature is probably linked to the diet or paramasticatory activity of at least one full population (Indonesian Homo erectus even if it is present on one African fossil). The interpretation and the use of Further research has to be done to determine where the preglenoid tubercle is present and also to better understand its functional significance in terms of its anatomy. the features is one of the big questions to which answers differ among the various anthropological schools, and which cannot be definitively answered here. Nevertheless, regardless of the theoretical approaches that lead people to accept one definition or another for a species and the taxonomic level at which the feature is used as a conclusion, I can propose my own scenario. Following the Hennigian principle, any feature can be used to make phylogeny. In this context, the preglenoid tubercle could be a feature that happened among some old African Homo erectus sensu lato (KNMER 3883), called Homo ergaster by authors (Andrews, 1984; Groves, 1989; Wood, 1992; Stringer, 1996) and that I considered as the type of another species (Zeitoun, 2000). The preglenoid tubercle still existed among Indonesian Homo erectus (Sangiran series) and also Archaic Homo sapiens or Homo soloensis (Ngandong-NgawiSambungmachan series). This interpretation suggests to consider the lack of the preglenoid tubercle among Sinanthropus as their belonging to another taxon. Thus, Sinanthropus were different from Indonesian taxa. More than merely shedding light onto what can be the differences between African Homo erectus sensu lato (Homo ergaster) and Asian Homo erectus sensu lato, the occurrence of the preglenoid tubercle triggers more definitive splitting of the taxon Homo erectus that, at least, appears to be paraphyletic. The history of the occurrence of the preglenoid tubercle may prove to be a key to understanding the links among Homo erectus sensu lato. Adopting other phylogenetic scholars’ points of view, the story can be different; but to conclude, the presence of a strong preglenoidal tubercle appears to be discriminating for some populations usually assigned to Homo erectus. ACKNOWLEDGMENTS I thank Professor T. Jacob, J. De Vos, J. Franzen, A. Langaney, M. Tranier, Dr. H. Widianto, and Mister Himawan for access to original material. I also thank J. Braga, M. Elyaqtine, P. Picq, S. Prat, P. Tassy, and A.M. Tillier for their contributions and suggestions; P. Janvier, E. Trinkaus, J. Dufour, M. Paalman, I. Tattersall, and three anonymous reviewers for their comments on the text. This work was supported by the Foundation Fyssen and the fellowship M. Bleustein-Blanchet. LITERATURE CITED Abe S, Ouchi, Y, Ide Y, Yonezu H. 1997. Perspectives on the role of the lateral pterygoid muscle and the sphenomandibular ligament in temporomandibular joint function. Cranio 15:203–207. Andrews P. 1984. An alternative interpretation of characters used to define Homo erectus. Courier Forschungsinst Senckenb 69:167–175. Anthony J. 1946. Le trou malaire des Atelidae et son intérêt dans la classification des singes platyrrhiniens. Mammalia 10: 69 –77. Barriel V. 1991. Caractères ostéologiques et odontologiques chez les Hominoidea (Primates, mammalia): Analyse de parcimonie. Bull Mém Soc Anthrop Paris 3: 45–72. 156 THE ANATOMICAL RECORD (PART B: NEW ANAT.) Barriel V. 1994. Les relations de parenté au sein des hominoidea et la place de Pan paniscus: Comparaison et analyse méthodologique des phylogénies morphologique et moléculaire. Thèse de doctorat de l’Université Paris 6 (inédit). Barriel V, Tassy P. 1991. La forme dans tous ses états (la morphologie et le binaire). Geobios 13:177–223. Bonde N. 1989. Erectus and neanderthalensis as species or subspecies of Homo with a model of speciation in hominids. In: Giacobini G, editor. Hominidae. Proceedings of 2nd International Congress of Human Paleontology. Milan: Jaca Book. p 205–208. Braga J. 1995. Variation squelettique et mesure de divergence chez les chimpanzés. Contribution de l’étude des caractères discrets. C R Acad Sci II 320:1025– 1030. Brauer E, Mbua, G. 1992. Homo erectus features used in cladistics and their variability in Asian and African hominids. J Hum Evol 22:79 –108. Burch J. 1970. Activity of the accessory ligaments of the temporomandibular joint. J Prosthet Dent 24:621–628. Campbell B. 1963. Quantitative taxonomy and human evolution. In: Washburn SL, editor. Classification and human evolution. Chicago: Aldine. p 50 –74. Couture C, Elyaqtine M, Houet F, Maureille B. 1990. Le crâne de l’abri Malaurie (Lot). Bull Soc Anthrop Sud-Ouest XXV 2:69 –81. Darlu P, Tassy P. 1993. La reconstruction phylogénétique. Concepts et méthodes. Collection biologie théorique. Paris: Masson. 273 p. Dubois E. 1940. The fossil human remains discovered in Java by G.H.R. von Koenigswald and attributed by him to Pithecanthropus erectus, in reality remains of Homo sapiens soloensis. Ned Akad Wetenschappen 43:841–854. Durband A. 2002. The view from down under: A test of the multiregional hypothesis of modern human origins using the basicranial evidence from Australasia. In: Acts of the XVIIth Congress of the Indo Pacific Prehistory association, Taipei. p 61. Falk D. 1986. Evolution of cranial blood drainage in Hominids: Enlarged occipital/marginal sinuses and emissary foramina. Am J Phys Anthrop 70:311–324. Griffin C, Hawthorn R, Harris R. 1975. Anatomy and histology of the human temporomandibular joint. Monogr Oral Sci 4:1–26. Groves CP. 1989. A theory of human and primate evolution. Oxford: Oxford University Press. 375 p. Hauser G, De Stefano GF. 1989. Epigenetic variants of the human skull. Stuttgart: Schwerzerbart. 301 p. Hublin JJ. 1986. Some comments on the diagnostic features of Homo erectus. Anthropos (Brno) 23:175–187. Hublin JJ. 1989. Les charactéres dérivés d’Homoerectus:relationavecl’augmentation de la masse squelettique. In: Giacobini G, editor. Hominidae. Proceedings of 2nd International Congress of Human Paleontology. Milan: Jaca Book. p 199 – 204. Jelinek J. 1981. Was Homo erectus already Homo sapiens? In: Ferembach D, editor. Les processus de l’hominisation. Paris: CNRS. p 85– 89. Kennedy GE. 1991. On the autapomorphic traits of Homo erectus. J Hum Evol 20: 355–371. Løvtrup S. 1987. On species and other taxa. Cladistics 3:157–177. Murray G, Phanachet I, Uchida S, Whittle T. 2001. The role of the human lateral pterygoid muscle in the control of horizontal jaw movements. J Orofac Pain 15: 279 –292. Msuya C, Harrisson T. 1996. The circum orbital foramina in Primates: A phylogenetic perspective. Kaupia 6:103–109. Nelson GJ. 1989. Species and taxa: Systematics and evolution. In: Otte D, Endler JA, editors. Speciation and its consequences. Sunderland, MA: Sinauer Associates. p 60 – 81. Nicolas A, Poirier P, Charpy A. 1926. Traité d’anatomie humaine. Vol. I. 4th ed. Paris: Masson. 667 p. Paturet G. 1951. Traité d’anatomie humaine. Vol. I. Paris: Masson. Picq P. 1983. L’articulation temporo-mandibulaire des Hominides fossiles: anatomie comparée, biomécanique, évolution, biométrie. PhD dissertation, University P. and M. Curie, Paris VI. Piette E. 1993. Anatomy of the human temporomandibular joint. An updated comprehensive review. Acta Stomatol Belg 90:103–127. Rightmire P. 1984. Comparisons of Homo erectus from Africa and South-East Asia. Courier Forschungsinst Senckenb 69:83– 99. Rightmire P. 1990. The evolution of Homo erectus. New York: Cambridge University Press. Rouvière H. 1954. Anatomie humaine, tête et cou. Vol. I. Paris: Masson. 544 p. Sato H, Strom D, Carlsson G. 1995. Controversies on anatomy and function of the ligaments associated with the temporomandibular joint: A literature survey. J Orofac Pain 9:308 –316. Sato H, Sindo K, Ezure H, Shimada K. 1996. Morphology of the lateral ligament in the human temporomandibular joint. FEATURE ARTICLE Oral Surg Oral Med Oral Pathol Oral Radiol Endod 8:151–156. Schmolke C. 1994. The relationship between the temporomandibular joint capsule, articular disc and jaw muscles. J Anat 184:335–345. Simpson GG. 1961. Principles of animal taxonomy. New York: Columbia University Press. 488 p. Strait DS, Grine FE, Moniz MA. 1997. A reappraisal of early hominid phylogeny. J Hum Evol 32:17–82. Stringer CB. 1984. The definition of Homo erectus and the existence of the species in Africa and Europe. Courier Forschungsinst Senckenb 69:131–143. Stringer C. 1987. A numerical cladistic analysis for the genus Homo. J Hum Evol 16:135–146. Stringer C. 1996. Current issues in modern human origins. In: Meikle W, Howell F, Jablonski N, editors. Contemporary issues in human evolution. San Francisco: California Acadamy of Science. p 115– 135. Tobias P. 1985. Single characters and total morphological pattern redefined the sorting effected by a selection of morphological features of the early hominids. In: Liss A, editor. Ancestors: The hard evidence. New York: E. Delson. p 94 –101. Trinkaus E. 1990. Cladistics and the hominid fossil record. Am J Phys Anthrop 83:1–11. Turner A, Chamberlain A. 1989. Speciation, morphological change and the status of African Homo erectus. J Hum Evol 18:115–130. Wiley E, Siegel-Causey D, Brooks D, Funk V. 1991. The compleat cladist a primer of phylogenetic procedures. Special publication n°19 of the University of Kansas, Museum of Natural History. Lawrence, KS: University of Kansas. 159 p. Wilkinson T. 1988. The relationship between the disk and the lateral pterygoid muscle in the human temporomandibular joint. J Prosthet Dent 60:715–724. Wood B. 1984. The origin of Homo erectus. Courier Forschungsinst Senckenb 69:389 – 406. Wood B. 1991. Koobi Fora research project. Hominid cranial remains. Oxford: Clarendon Press. Wood B. 1992. Early hominid species and speciation. J Hum Evol 22:351–365. Wood B. 1994. The oldest hominid yet. Nature 371:280 –281. Zeitoun V. 1996. Cladistique et paléoanthropologie: le cas Homo erectus (Dubois, 1894). PhD dissertation, University of Bordeaux I. Zeitoun V. 2000. Révision de l’espèce Homo erectus (Dubois, 1893). Bull Mem Soc Anthropol Paris 12:1–200.