ARTICLE

Microscopic Investigation in Fossil Hominoidea:A Clue to Taxonomy, Functional Anatomy, andthe History of DiseasesMICHAEL SCHULTZ*,†

Ten selected samples of fossilized bones (including Australopithecus, Homo erectus, Homo neandertalensis , andHomo sapiens sapiens) were examined by light microscopy using plane and polarized light. The histomorphologicalfindings show that microscopic research adds much to what can be ascertained by marcoscopic examination or byX-ray techniques. In particular, emphasis was placed on taxonomy, functional anatomy of bones, and causes of someof the diseases of early hominids. Anat Rec (New Anat): 257:225–232, 1999. r 1999 Wiley-Liss, Inc.

KEY WORDS: paleoanthropology; paleopathology; microscopic anatomy; comparative anatomy; human fossil; histology; lightmicroscopy; dinosaur

Light microscopy—particularly withpolarized light—microradiography,and scanning-electron microscopy arevaluable tools for paleoanthropolo-gists, not only for comparative ana-tomic studies but also for differentialdiagnoses of dry bones and fossils.Bone changes in prehistoric and his-toric skeletal remains, which cannotbe differentiated by macroscopicanalysis, are easily revealed by micro-scopic techniques.12,13,15 Thus, alter-ations caused intra vitam by disease,can clearly be differentiated fromchanges due to postmortem reac-tions.12 The nature of bone tumorsand tumorous lesions can be detectedas well as the different structures oc-curring through specific and non-

specific bone inflammations.2,20 Also,changes caused by metabolic diseases,processes of aging, decrease or in-crease of functional activity, etc., areidentifiable with the aid of micro-scopic methods.12

Until now, these methods have neverbeen used in an extended and com-parative study using the remains offossil hominids and other Hominoi-dea.1 The reasons are, of course, wellknown: the technique which is used toobtain bone samples is invasive. Thismeans that the fossil must be partlydamaged by removal of a small bonesample. However, since the samplecan be replaced by a cast, the morphol-ogy and the basic information of sucha fossil will ultimately not be dis-turbed. We should remember that re-cent and fossilized bones are archivesof life that preserve important informa-tion. As a rule, microanalysis of fossilsprovides insights into the past lives ofour recent and earliest ancestors thatcannot be gained by any other tech-nique.

It is well known that archeological,i.e., subfossilized and fossilized, boneis frequently affected in the ground byvarious factors, e.g. roots of plants,fungi, algae, bacteria, protozoa, in-sects and their larvae, worms, andmechanical agents such as water andcrystals.12,16 Unfortunately, even to-

day, little is known about the physiol-ogy of the flora and fauna of cadaverspreserved over many centuries or thou-sands of years in the ground. All thesepostmortem factors produce damagethat can falsely be diagnosed as le-sions caused intra vitam by diseases.Therefore, knowledge of decomposi-tion and diagenesis is very importantin diagnosing fossilized bone by micro-scopic techniques.

This paper aims to demonstrate thenecessity of microscopic research infossilized human bones. As there arenow several reliable techniques avail-able13,18 which work with relativelysmall samples, paleoanthropologistswould be well-advised to use theseuseful tools for their research. Particu-larly, the examination of thin-groundsections viewed in polarized light yieldthe most reliable results.13 Therefore,this report aims to demonstrate thelarge amount of information that mi-croscopic research of fossilized bonescan contribute to our knowledge of tax-onomy, functional anatomy, and thecauses of diseases of early hominoidea.

THE STUDYFor this study, 10 different fossil speci-mens were used. Sample 1: piece ofthe middle part of the shaft of a hu-merus of an Australopithecus fromSwartkrans (South Africa), dated at

Dr. Schultz is Professor of Anatomy atthe Center of Anatomy, Go ¨ ttingen Uni-versity. His scientific research and teach-ing interests cover human anatomy, physi-cal anthropology, primatology, andpaleopathology. He is President of the Ger-man speaking Anthropologists (GfA) andone of the Managing Editors of HOMO.

†Dedicated to Professor Dr.med. Hans-Jurg Kuhn on the occasion of his 65 th

birthday.*Correspondence to: Prof. Dr.med. Dr-

.phil.nat. Michael Schultz, ZentrumAnatomie der Georg-August-Universi-tat, Kreuzbergring 36, D-37075 Go ¨ t-tingen, Germany. Fax: 0049–551–39–7028; E-mail: mschult1@gwdg.de

THE ANATOMICAL RECORD (NEW ANAT.) 257:225–232, 1999

r 1999 Wiley-Liss, Inc.

ca. 1–2 my; Sample 2: small fragmentsof a long bone of an Australopithecusfrom Sterkfontein (South Africa), ca.2.2–3 my; Sample 3: fragment of theparietal bone of the Homo erectus(O.H. 9) from Olduvai Gorge (Tanza-nia), ca. 1.1 my; Sample 4: small frag-ment of the greater wing of the sphe-noid bone of Homo erectus fromPetralona (Greece), ca. 300,000 years;Sample 5: right and left ulnae (a,b) ofthe Homo neanderthalensis found in1856 near Dusseldorf (Germany), ca.65,000 years; Sample 6: fragment of aclavicle of an individual of Homo nean-derthalensis from Krapina (Croatia),ca. 70,000 years; Sample 7: small frag-ment of the skull vault (probably pari-etal bone) of an individual of Homoneanderthalensis from Vindija (Croatia),ca. 35,000 years; Sample 8: piece ofthe right part of the occipital boneclose to the occipitomastoid suture ofthe female Homo sapiens from Kelster-bach (Germany) that is one of theoldest examples of anatomically mod-ern man in Central Europe (R. R. R.Protsch), ca. 32,000 years; Sample 9:transverse section of the femur of anadult Pan paniscus from the Depart-ment of Anthropology, Arizona StateUniversity at Tempe (USA), ca. 30 y,and for comparison purposes; Sample10: a piece of a vertebra of a dinosaurwith the oldest tumor found to date,representing a hemangioma (Arizona),ca. 140–150 my.

From all these samples, thin groundsections were produced at a thicknessof 50 and 70 µm using the techniquespreviously described.2,18 The sectionswere viewed by light microscopes(Zeiss ‘‘Ultraphot II’’, Leica ‘‘LEICADMRXP’’) in normal (plane) and polar-ized light using an additional hilfsob-ject red 1st order (quartz) as compensa-tor.

LIGHT MICROSCOPY,DIAGENESIS, AND FOSSILSMicroscopic research on fossilizedbones is an interdisciplinary methodthat comprises anatomy, anthropol-ogy, and paleontology. Light micros-copy, particularly using polarized light,yields more diverse information abouta sample than scanning electron-mi-croscopy, which can help in diagnos-ing external structure but gives noinformation about the internal mor-

phology.6 For light microscopy, thinground sections have to be prepared.As a rule, it is difficult to produce thinground sections from fossilized bonesamples because the samples arebrittle, hard, and always extremelydifficult to handle. Although Moodie,9

as one of the first in his field, workedwith ground sections of fossilizedbones (e.g., dinosaur), this kind ofresearch has never become very popu-lar. The reasons are probably the diffi-culties in producing such sections andthe great amount of experience that isnecessary to diagnose fossilized andsubfossilized bones.

Sometimes, fossilized bone cannotbe investigated microscopically be-cause of the poor state of preservation.As a rule, however, fossilized bone ismuch better preserved than prehis-toric or historic bones. This soundsparadoxical, but is easy to understand.Of course, the overwhelming majorityof the bones of early hominids havebeen completely destroyed by diagen-esis (the changes that occur normallywithin sediment during and after itslithification).16 Only a very small num-ber of bones or bone fragments were,relatively immediately, embedded byprotective soil (e.g., in caves) after thedeath of an individual.21 Such bonetissue was well preserved. The preser-vation of the piece of the occipitalbone of the female skull of Homosapiens from Kelsterbach is excel-lently preserved. The ground section(sample 8) presents structures as clearas in fresh bone (Fig. 1). No diseasescould be diagnosed from analysis ofthis sample. Also extremely well pre-served is the small bone fragment ofthe greater wing (sample 4) of theskull from Petralona representingHomo erectus (Fig. 2). Again, in this

sample there are no vestiges of patho-logical alterations. A small fragmentof the skull vault (parietal bon) of aNeanderthal individual from Vindjia(sample 7) is relatively poorly pre-served, but organic material (colla-gen) is still visible in this ground sec-tion (Fig. 3). The texture of the calcifiedbone tissue is partially destroyed bymicrofractures. Nevertheless, no ves-tiges of diseases could be diagnosed.

It is well known that lamellar bonemay be compact or cancellous. Theexamples presented in this study aremainly composed of compact lamellarbone. The microstructure of lamellarbone, especially the course and theorientation of collagen fibers whichare birefringent, can be easily ob-served in polarized light.17 Fossilizedbones can also be examined by thismethod if the matrix is relatively wellpreserved. The micromorphology ofcompact bones in various mammalianspecies is varied (e.g., 3). Therefore, itis possible to differentiate betweenhuman and other animal bones bymicroscopic techniques.

THE MICROSTRUCTURE OF BONEIN AUSTRALOPITHECUSThe samples of the two Australopithe-cus specimens are preserved to differ-ent extents. The sample from Swart-krans (sample 1) is a short butcomplete piece of the diaphysis of thehumerus and is very well preserved,but the smaller piece from Sterkfon-tein (sample 2) is only a fragment ofthe wall of a long bone diaphysis thatis only moderately well preserved. Thecompact bone substance of the twosamples is characterized by relativelysmall Haversian canals which have aminimal diameter of approximately12 µm and a maximal diameter ofapproximately 74 µm (mean value 30µm, median value 31 µm). As thewhole transverse section of the middlepart of the shaft of the humerus of theAustralopithecus from Swartkransshows (Fig. 4), the external basic la-mella is extremely thick, but very welllamellated. The measurements of theground section yield minimum thick-ness of 913 µm and a maximum thick-ness of 1509 µm. Thus, the externalbasic lamella takes up approximatelyone-third of the diameter of the wall ofthe diaphysis of the humerus. The

As a rule, microanalysisof fossils provides insightsinto the past lives of our

recent and earliestancestors that cannot be

gained by any othertechnique.

226 THE ANATOMICAL RECORD (NEW ANAT.) ARTICLE

Figure 1. Kelsterbach hominid. Transverse section through externallamina of the occipital bone (sample 8).Thin ground section (50 µm).Microscopic view using polarized light and hilfsobject red 1st order(quartz) as compensator. Magnification 253.

Figure 2. Petralona skull. Transverse section through fragment of thegreater wing of the sphenoid. Bone trabecula and sinter (sample 4).Thin ground section (50 µm). Microscopic view using polarized light andhilfsobject red 1st order (quartz) as compensator. Magnification 1003.

Figure 3. Neanderthal man from Vindija. Transverse section throughexternal lamina of the skull vault, probably parietal bone (sample 7)demonstrating postmortem microfracture. Thin ground section (50µm). Microscopic view using polarized light and hilfsobject red 1st

order (quartz) as compensator. Magnification 253.

Figure 4. Australopithecus from Swartkrans. Transverse sectionthrough humerus (sample 1). External basic lamella, compact bonesubstance, internal basic lamella, and secondary fill of medullarycavity. Thin ground section (50 µm). Microscopic view using polarized lightand hilfsobject red 1st order (quartz) as compensator. Magnification 253.

Figure 5. Australopithecus from Swartkrans. Osteons in transversesection through humerus (sample 1). Small Haversian canals. Thinground section (50 µm). Microscopic view using polarized light andhilfsobject red 1st order (quartz) as compensator. Magnification 1003.

Figure 6. Pan paniscus. Femur (sample 9) Transverse section throughfemur (sample 1). Endosteal area of compact bone substance. Thinground section (50 µm). Microscopic view using polarized light andhilfsobject red 1st order (quartz) as compensator. Magnification 253.

Figure 7. Dinosaur bone. Transverse section through a piece of thevertebral body (sample 10). Bone trabeculae in area of hemangi-oma. Medullary cavities filled by products of fossilization process. Thinground section (50 µm). Microscopic view using polarized light andhilfsobject red 1st order (quartz) as compensator. Magnification 253.

Figure 8. Homo erectus from Olduvai Gorge (O.H. 9). Transversesection through external lamina of the occipital bone (sample 3).Thinground section (50 µm). Microscopic view using polarized light andhilfsobject red 1st order (quartz) as compensator. Magnification 253.

internal basic lamella, however, is com-paratively thin (75 µm). The compactzone of the transverse section, whichis found between the external and theinternal basic lamella and which isbuilt up of Haversian systems andinterstitial lamellae, measures between2071 and 2796 µm.

There is no apparent difference inthe structures of compact bone ofanatomically modern man and Nean-derthal man. However, it is strikingthat the compact bone of the shaft ofthe humerus of the Australopithecusspecies from Swartkrans (Fig. 4) showsstructures significantly different fromrecent humans and Neanderthal man.These different features are the thick-ness of the external basic lamella andthe size of the Haversian canals (Fig.5). The external basic lamella of Austra-lopithecus is very well lamellated, butextraordinarily thick. It takes up ap-proximately one-third of the diameterof the wall of the diaphysis of thehumerus. In anatomically modern manand Neanderthal the external basiclamella represents in the transversesection—if present—only a very finerim. It should be kept in mind that thethickness and the presence of the exter-nal basic lamella depend on age, i.e.,in modern man, the external basiclamella is completely resorbed, as arule, in individuals who are older thanapproximately 40 years. Thus, it couldbe assumed that the individual fromSwartkrans was probably relativelyyoung at the time of death. The secondfeature of interest, which is differentin Australopithecus, is the relativelysmall size of the Haversian canals.This could be observed in the speci-mens from Swartkrans and Sterkfon-tein. It is remarkable that the luminaof the Haversian canals in Australopi-thecus (Figs. 4,5) have more or less thesame size as in the chimpanzee Panpaniscus (Fig. 6).

MICROSCOPY YIELDS CLUES TOTAXONOMYThe ground section of the chimpanzee(sample 9) from Arizona Zoo is verywell preserved. The Haversian canalsare relatively small which is character-istic of Pan species (Fig. 6). The diam-eters of the canals are in the samerange as those of the canals observedin Australopithecus. From the micro-

scopic point of view, the basic struc-ture of the compact bone substance,i.e., the Haversian systems, is verysmilar in Australopithecus and Panand different from those in Homo.Thus, the Australopithecus speciesfrom Swartkrans and Sterkfontein areprobably more closely related to Panpaniscus than to Homo neandertalien-sis or Homo sapiens. There is no histo-morphological difference between thelatter two. These results demonstratethat there may be taxonomic differ-ences detectable at the microscopiclevel. This means that the microscopicinvestigation could help in the futurein the determination of the taxonomyof fossilized bones of different homini-dea, particularly, when only a few smallbone fragments are preserved.

ANALYSIS OF THE OLDEST TUMORBY LIGHT MICROSCOPYTo date, little has been reported on thepaleohistopathology of fossilized hu-man bones, probably on the basis ofthe hypothesis that no significant mi-croscopic structures are preservedwhich could make reliable diagnosespossible. However, it was recently dem-onstrated10 that even in an Upper Ju-rassic fossil of a dinosaur, a tumor(hemangioma) could be diagnosed bymicroscopic techniques (Fig. 7). Forcomparison purposes, this oldestknown tumor examined in the verte-bra of a dinosaur10 is presented todemonstrate that also in very old fos-sils the diagnosis of diseases is pos-sible (sample 10). In the ground sec-tion, there are coarse bone trabeculaein a net-like structure which are situ-ated between enlarged spaces repre-senting cavities characteristic for hem-angiomas.

THICKENED SKULL VAULTSCAUSED BY PATHOLOGICALPROCESSES?The sample from the parietal bone(sample 3) of the Homo erectus fromOlduvai Gorge (O.H. 9) is extraordinar-ily well preserved.8 All structures areregular. The well lamellated externallamina is relatively thick (Fig. 8). Su-perficially observed, there is no appar-ent evidence of pathological changes.Of course, vestiges of pathological pro-cesses are relatively rare in fossilizedbones. Sometimes, there are changesthat cannot be diagnosed convinc-ingly, caused by pathological agents.For instance, there are different hy-potheses on the phenomenon of therelatively thick skull vault of Homoerectus. In prehistoric skulls, a thick-ened skull vault is frequently due todiseases15 such as a healed inflamma-tory process (e.g., osteomyelitis) orsome deficiency diseases (e.g., ane-mia, rickets, scurvy). Particularly, inchronic anemia a pronounced enlarge-ment of the diploe is observed which isresponsible for the thickening of thebone. As a rule, vestiges of these dis-eases are found on the external part ofthe parietal and the frontal bones. Theresult of the microscopic examinationof a fragment of the parietal bone ofthe Homo erectus from Olduvai Gorge,which is called O.H. 9, demonstratesthat there are no vestiges of diseases.The external lamina of the skull boneis, as a rule, much thicker than inrecent man, but resembles the struc-ture seen in early neolithic popula-tions called the bandkeramiker. Up tonow, the causes for the enlargement ofthe external lamina are still unknown.Nevertheless, there is one hypothesisthat this thickening of the skull vault isdue to hypervitaminosis A14 that couldtheoretically be induced by extensiveeating of animal liver, which seemshardly probable.

SOCIAL CARE IN NEANDERTHALMANThe skeleton of the Neanderthal manfound in the Kleine Feldhofer Grottein 1856 near Dusseldorf, Germany, wasrecently re-examinedbyan interdisciplin-ary group.7,11 Bone samples were takenfor the microscopic analysis.19 Thepreservation of the compact bone

Paleopathology,particularly microscopicresearch, enlarges our

knowledge of theetiology and theepidemiology ofdiseases from the

beginning of mankind.

ARTICLE THE ANATOMICAL RECORD (NEW ANAT.) 229

Figure 9. Neanderthal man from the Nean-der Valley. Transverse section through leftulna (sample 5b). Compact bone demon-strating osteoporosis. Thin ground section (50µm). Microscopic view in plane (normal) light.Magnification 253.

Figure 10. Neanderthal man from the Nean-der Valley. Transverse section through rightulna (sample 5a). Compact bone. Thinground section (50 µm). Microscopic view inplane (normal) light. Magnification 253.

Figure 11. Neanderthal man from Krapina.Transverse section through fragment of aclavicle (sample 6). Osteoporosis. Thin groundsection (50 µm). Microscopic view in plane(normal) light. Magnification 253.

substance varies from fairly good tovery good. The individual’s age checkedby the method of Kerley4 and Kerleyand Ubelaker5 suggests the mature agegroup.

In the left ulna which had beenfractured during the Neanderthal’s life-time (sample 5a), enlarged Haversiancanals and an increase of spongy bonein the endosteal area of the primarilycompact bone substance demonstrateevidence of osteoporosis which can-not be the result of the age-dependentbone involution (Fig. 9). The bonesubstance of the right ulna (sample5b) is regular and shows no vestiges ofosteoporosis (Fig. 10).

This mature male Neanderthal mansuffered from a fracture of the leftulna which healed during his lifetime.Using light microscopy, possiblechanges in the micromorphology ofthe affected bone which could be dueto the trauma were sought. The resultsare indeed striking. In comparison tothe right ulna (Fig. 10), the left showssevere osteoporosis in the fracturedbone (Fig. 9), likely caused by inactiv-ity atrophy.12 Thus, after the traumawhich broke the left ulna, this indi-vidual probably could not use his leftarm for the rest of his life, and so wasunable to do all the necessary things ofevery day life in the right way. Thisobservation is important, because itsuggests strongly that there was somekind of social care in the times ofNeanderthal man.

OLD AGE OSTEOPOROSIS INNEANDERTHAL MANAmong the samples studied, there isanother case of osteoporosis in Nean-derthal man in the form of generalizedenlargement of blood vessel canals.The clavicle of one of the individualsfrom Krapina (sample 6) shows dis-tinct features of this disease (Fig. 11).The preservation of the organic mate-rial (collagen) in the sample takenfrom the clavicle of a Neanderthalindividual from Krapina is not verygood. The compact bone substance ofthe shaft of the clavicle is character-ized by enlarged Haversian and non-Haversian canals. The external sur-face of the fragment of this bone issmooth. Other changes are not observ-able. The cause of this porosity cannotbe established. Old age osteoporosity

is probably the most reliable diagno-sis.

PERSPECTIVESThe results of the microscopic exami-nation of samples of fossilized bonesdemonstrate that this technique makesit possible to gain reliable informationon not only micromorphology but tax-onomy and histopathology as well.Thus, valuable data on cladistic rela-tionships, functional anatomy, and thediseases of ancient man can be gath-ered and analyzed. In future work, it isimportant to keep in mind that thehistomorphologic analysis of fossil-ized human bones is a vital necessity.Paleopathology, particularly micro-scopic research, enlarges our knowl-edge of the etiology and the epidemiol-ogy of diseases from the beginning ofmankind. Thus, the history of diseasescould be better investigated if humanfossils were examined by microscopictechniques.

ACKNOWLEDGMENTSThe author sincerely thanks Prof. Dr.Dr. R.R.R. Protsch, Frankfurt Univer-sity (Germany), Prof. Dr. W.E. Maier,Tubingen University, Dr. A. Nkini, Da-ressalam (Tanzania), Prof. Dr. N. Xiroti-ris, Komotini University (Greece), Dr.H.-E. Joachim and Dr. R. Schmitz,Rheinisches Landesmuseum Bonn(Germany), Prof. Dr. C.F. Merbs, Ari-zona State University, Prof. Dr. B.M.Rothschild, The Arthritis Center ofNortheastern Ohio for the bonesamples, M. Brandt, I. Hettwer-Steeger,Th. Schlomm, and T. Walde, Got-tingen University for technical assis-tance, and C. Maelicke, Gottingen Uni-versity for reading the English text.

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