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Mandibular Fossa of Homo Erectus

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Mandibular Fossa of Homo Erectus THEANATOMICALRECORD266:142–145(2002) DOI10.1002/ar.10053 BoneBits MandibularFossaofHomoerectus: AResponsetoDurband 1,2 3–6 4–7 KENMOWBRAY, *SAMUELMA´ RQUEZ, ANDERICDELSON 1DivisionofAnthropology,AmericanMuseumofNaturalHistory, NewYork,NewYork 2DepartmentofAnthropology,RutgersUniversity,NewBrunswick,NewJersey 3CenterofAnatomyandFunctionalMorphology,MountSinaiSchoolofMedicine, NewYork,NewYork 4DepartmentofAnthropology,LehmanCollege/CUNY,Bronx,NewYork 5Ph.D.PrograminAnthropology,CUNYGraduateCenter,NewYork,NewYork 6NewYorkConsortiuminEvolutionaryPrimatology,CityUniversityofNewYork, NewYork,NewYork 7DivisionofPaleontology,AmericanMuseumofNaturalHistory, NewYork,NewYork Durband(2002)hascriticizedourrecentanalysisof WT15000alsoshowsaflatandlongfossa,butthe Sambungmacan3(Delsonetal.,2001)withregardtothe squamotympanicfissurecoursesinitsdeepestpor- implicationsofthesquamotympanicfissure(SQF)andthe tion.However,itisnotedagainthatgrowthhasnot postglenoidtubercle(PGT)forgroupingHomoerectusfos- beencompletedinthisspecimen(seeabove).Finally, sils.Hiscriticismisbasedupontwoabstractsandan OH9presentsamandibularfossathatissimilarboth unpublished masters thesis (Durband 1997, 1998; initsshapeandinthepositionofthesquamotym- DurbandandKidder,2000)anddependsoninterpretation panicfissuretotheNgandongandotherIndonesian oftheobservedmorphologyaswellasthetaxonomic specimens.ItappearsthatthistraitunitestheIndo- “weight”ofthesecharacters.Herewetaketheopportunity nesianH.erectusspecimenstotheexclusionofthe tobrieflyrespondtohisqueriesandprovideobservations otherAsianandAfricanones,withtheexceptionof thataresomewhatmoreextensive. OH9andpossiblyalsoWT15000....” Delsonetal.(2001:390–392)wrote: Durband(2002)essentiallyarguesthatwithintheman- dibularfossa,thefeature“courseoftheSQF”mustbe “Sm3alsoshareswiththeNgandongspecimens furtheratomizedintovariousstatesintermsofitsorien- thedistinctmorphologyandrelativepositionofthe tation,depth,andpositionwithrespecttocontiguous squamotympanicfissure(Fig.7).Inhisdescriptionof structures,especiallythePGT.Hefurtherquestionsthe theNgandongcrania,Weidenreich(1951,p.273–4) presenceofaPGTinavarietyofH.erectusspecimens.In notedboththedeepandanteroposteriorlyshortshape fact,Durbanddefinesa“Ngandonggroup”(whichforhim ofthemandibularfossaandthecourseofthesqua- includestheSambungmacan1and3andNgawifossils) motympanicfissure(whichhemistakenlytermedthe basedonthesharedpossessionofcharacterstatesofthe “Glaserian”fissure;seeMa´rquezandMowbray,in resultingsubdividedfeatures.Thus,inhisopinion,spec- preparation)asbeingcharacteristicforthisgroup.He imensoftheNgandonggrouppresentanSQFwhichlies observedthatintheNgandongspecimensandinSan- entirelyinthedeepestpartofthemandibularfossabut giran4thesquamotympanicfissurecoursesmedio- lackaPGT,whileinotherspecimensthelateralendofthe laterallyandcoincideswiththedeepestportionofthe fissurecurvesaroundthePGTanddepartsfromthedeep- mandibularfossa,whereasintheChineseH.erectus estportionofthefossa. specimensandinmodernhumansthisfissureispo- OurpublishedtextfollowedWeidenreich(1951,ascited sitionedmoreposteriorly,andthefloorofthefossa above)inexaminingonlythefirstofthesefeatures.Wedid descendstowardtheposteriorwallbeforereaching notoriginallymakeextensivecomments,asourgoalwas thefissure.Inthelastfiftyyears,manyadditionalH. tolistthepresenceorabsenceofcharactersattributedto erectuscraniapreservingthisregionhavebeenrecov- ered.TheIndonesianspecimensSangiran17andSm 1showamorphologywhichissimilartothatpre- servedintheNgandonggroup.TheAfricanrepresen- *Correspondenceto:KenMowbray,DivisionofAnthropology, tativesaredividedintheexpressionofthisfeature. AmericanMuseumofNaturalHistory,NewYork,NY10024.Fax: ER3733and3883haveaflatandanteroposteriorly (212)769-5334.E-mail:[email protected] longmandibularfossaandasquamotympanicfissure Received30October2001;Accepted7November2001 thatrunsposteriortothedeepestportionofthefossa. Publishedonline13February2002 ©2002WILEY-LISS,INC. MANDIBULAR FOSSA OF H. ERECTUS: A RESPONSE 143 Fig. 1. a: Right mandibular fossa in basal view. Note the division of the SQF medially into the petrotym- panic fissure posteriorly (arrowheads) and petrosquamous fissure anteriorly (arrows), caused by the en- croachment of the tegmen tympani into the mandibular fossa. b: Left mandibular fossa in basal view. Note the bilateral difference in the course of the SQF. © Mowbray and Ma´ rquez. H. erectus (s.l.1) by a number of sources, present it in In humans, the mandibular fossa derives from two mes- tabular form, and attempt to relate Sm 3 to other speci- enchymal condensations—a glenoid blastema associated mens. Our aim was to extend Weidenreich’s data by ex- with the temporal bone, and a condylar blastema associ- amining casts of specimens unknown in the 1940s and ated with the mandible—that slowly fuse together around briefly report our assessments. Moreover, we offered no the 7th to 12th week in utero. At 23 weeks, the petrous new information on Sangiran (Sgr) 4 in the quoted excerpt portion and tympanic ring begin synostosis, producing the but merely cited Weidenreich’s observation, a point petrotympanic fissure through which pass the chorda tym- Durband apparently misunderstood. pani nerve and residual tissue of the malleolar disc liga- ment (Sperber, 2001). At birth, the tympanic ring fuses MANDIBULAR FOSSA with the squamous part of the temporal bone, forming the In order to respond to Durband’s assessment of features SQF. The tympanic ring continues to grow laterally (in within the mandibular fossa, we find it is necessary to response to the lateral direction of growth of the neuro- accurately describe the ontogeny of this basicranial por- cranium) forming the tympanic plate. Often, the thinned tion in anatomical position as a baseline for understand- anterior surface of the petrous portion of the temporal ing the topography involved in the course of the SQF. bone—or the roof of the tympanic cavity (i.e., the tegmen tympani), encroaches into the SQF and later ossifies, thus further dividing this region into the petrotympanic fissure posteriorly and the petrosquamous fissure anteriorly (Fig. 1a). 1 Some authors have placed these and other African fossils in H. As the mandibular fossa develops, it increases in size ergaster (Tattersall, 1995; Wood, 1991) or even H. leakeyi (Clarke, and changes in shape due to epigenetic constraints (e.g., 1990), but for the present we retain a broad definition of H. erectus, which subsumes all of these specimens. We note that at strain generated during chewing episodes). For example, least one (D 2282) of the recently described crania from Dmanisi, in living humans the fossa is generally flat and wide Georgia, which have been suggested to be most similar to African anteroposteriorly, as it is in chimpanzees, gorillas, austra- “H. ergaster” (Gabunia et al., 2000; Balter and Gibbons, 2000), in lopiths, and early Homo.InH. erectus (s.l.), however, the fact has facial features strongly reminiscent of Sgr 17. mandibular fossa is typically much narrower anteroposte- 144 MOWBRAY ET AL. riorly and higher from the base of the tympanum to the cause of age differences, sexual dimorphism, and morpho- roof. In some circumstances, it can appear cone-shaped, as logical variability generated through differences in diet. seen in Sm 3 and Ng 11. Moreover, the SQF is often bilaterally dissimilar within As the fossa increases in size ontogenetically, the pos- individuals (compare Fig. 1a and b), especially in seden- terolateral edge of the mandibular roof (the anterior edge tary modern populations that tend to exhibit malocclusion of the SQF) may develop an eminence that is generally disorders and periodontal disease. called either a postglenoid process or a PGT. The PGT is When dealing with fossil humans, it is rare to find simply a build-up of bone anterior to the tympanic plate bilaterally well-preserved (i.e., unbroken) mandibular fos- that serves as an attachment site for ligaments of the sae, which adds to the confusion of describing this region superior lamina—a band of elastic fibers that allows the at such an atomized level of inspection. In addition, the disc between the temporal bone and the mandibular con- highest point of the mandibular fossa can change depend- dyle to translate forward. The area of temporal bone ing on the orientation of the skull in the sagittal plane. One where a PGT has the potential to develop also marks the might consider orienting in the three-point plane configura- most lateral margin of the mandibular joint capsule. The tion of the Frankfurt Horizontal (FH), but since many of the size and shape of the resulting eminence will determine fossil specimens lack faces, the FH is difficult to accurately its descriptive epithet. Unfortunately, there is no clear establish. Considering these cautionary notes, we now ad- distinction among the use of such terms as “postglenoid dress the specific comments offered by Durband (2002) con- process” and “tubercle” since much of the description of cerning the feature “course of the SQF.” the PGT is highly subjective and somewhat arbitrary due In the Indonesian fossils, we found that Sgr 17, Sm 1, to its overall continuously varying morphology. Nonethe- and Sm 3 all are similar to the Ngandong series in that less, if present, the PGT (or process or spine) can develop their SQF lies in the highest portion of the mandibular in modern humans as early as the first year of life fossa. We do not agree that Durband’s Figure 1 (modified (Scheuer and Black, 2001) and can become extensively after Wood, 1991, by the addition of the position of the altered
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