Osteometric Variation of the Human Spine in Central Europe by Historic Time Period and Its Microevolutionary Implications

\l N/

PhD - Thesis

by osteometric variation of the Human spine in central Europe

Historic Time Period and Its Microevolutionary Implications

Dr. med. Frank Jakobus Rühli

Unit Biological Anthiopology and Comparative Anatomy Research

Department of Anatomical Sciences

The University of Adelaide - Australia

Supervisor:

Prof. Dr. M. Henneberg

of Adelaide - Australia Department of Anatomical sciences, The university

Co-SuPervisor:

Prof. Dr. Dr. M. Schultz

ZentrumAnatomie,Georg-August-UniversitätGöttingen-Deutschland

June 2003 Table of Contents

Tables ..... 5

Figures ... 7

Abstract .. 9

Statement 11

Acknowledgments .. t2

Introduction I4

Anatomical aspects of the human spine ...'...... ' l4

sexual dimorphism and age-related adaptalions of the human spine .... 27

Biomechanics and spinal morphology...... '.'.' 31

Osteometric findings of earlier spinal studies ... 34

lmpact of osteometric research of the human spine ...... ' 51

Spinal pathologies - especially lower back pain 59

Clinical and dynamic assessment of the spinal neural pathways""' 68

Historical perspectives of spinal disorders and morphometry "'..". 78

The non-human spine 83

Major evolution of the human spine and its physiological adaptations 87

Anatomical variations of the spine ...... 92

Microevolutionary and secular trends in human anatomy ...'. 97

Aim of the study ...... 105

Hypothesis to be tested .. 105

Material 106

Supplementary information on the samples ... rt2

2 F. J. Rühli - Osteometric Variation of the Human Spine Methods 119

Measurements ...... 119

Technical equipment t28

Estimation of intra- and inter-observer error r29

Examination permission 135

Data analysis 135

Critical sample size ...... 136

Modern samples ... r37

Results t39

Sex and age composition of the samples 139

Descriptive statistics of the measurements ..'..'...' t39

Osteometric data of the whole sample t42

Sexual dimorphism 150

Side differences of spinal measurements ..'..'.....'...... " "' 151

Inter-correlations of all measurements 151

Correlation of examined variables with individual age "' t52

Microevolutionary trends since the Late Pleistocene ""' 158

Secular changes of the intervertebral foramen in the modern samples .....'. 163

Analysis of variance: variable means with respect to time before present ...... '.'...... 166

Analysis of variance: variable standard deviations with respect to time before presentlT2

Principal components analysis of the spinal variables """""""176

Discussion 178

Osteometric knowledge of historic spines 178

Study limitations 179

J F. J. Rühti - Osteometric Variation of the Human Spine Comparative analysis of the results ...... 187

Variation of spinal morphometry due to sex and age 225

Inter-populational variations of spinal morphometry 230

Relation of geography and society to spinal morphometry ..'.'...... 230

Influence of stature and body size on spinal morphometry 234

Microevolutionary trends in body size and robusticity ."."-..'..236

Alterations of brain and skull morphometry as models for the spinal microevolution 256

Microevolution and secular trends of the spine and their possible etiologies ...... 258

Importance and functional implications of osteometric spinal data 273

Osteometric dimensions and their possible clinical value: the intervertebral foramen 275

Conclusions 278

References 280

Appendix 334

F. J. Rühli - Osteometric Variation of the Human Spine 4 Tables

Table 1: Osteometric studies of recent and historic spine samples 53

Table2: Morphological studies of spines in living people and cadavers ...... 55

Table 3: Individuals / samples included in the present study 108

Table 4 Measurements used t23

Table 5: Inter-observer erïor of selected vertebral and long bone dimensions ...... 134

Table 6: Composition of modern samples St. Johann and "Geneva", with individually

known sex and age 138

Table 7: Mean, standard deviation (SD) and number of individuals measured (N) of

all variables for whole sample and modern subgroups '..'...... I44

Table 8: Pearson correlation coefficients (r) of caudal intervertebral foramen width

with birth year ..... 164

Table 9: Pearson correlation coeff,rcients (r) of intervertebral foramen height with birth

year...... '.".... 165

Table 10: Inter-correlation of anterior vertebral body height at various levels ...... 186

Table I 1: Vertebral body height (mm) of various samples ...... '.. 188

Table 12: Vertebral body diameters (mm) of various samples ..'...... 196

Table 13: Maximum pedicle height (mm) of various samples .. 202

Table 14: Neural canal sizes (mm) of various samples 207

Table 15: Intervertebral foramen width (mm) of various samples ...... '...... '.'.215

Table 16: Correlation coefficient between individual stature and foramen magnum

breadth 2t7

Table 17: Foramen magnum dimensions (mm) of various samples 2t8

F. J. Rühl¡ - Osteometric Variation of the Human Spine 5 Table 18: Length of spinous process as a percentage of the sagittal diameter of the

vertebral body ...... 221

Table 19: Transverse process width (mm) of various samples 222

Table 20: Maximum humeral length (mm) of various samples ......

Table 2l: Femur length (mm) and femur head breadth (mm) of various samples 240

Table22: Femur mid-shaft circumference (mm) of various European samples .. .241

Table 23: Bi-iliac breadth (mm) of various European samples ..... 242

Table24 Estimated statures (cm) of various European samples .. ..243

F. J. Rühli - Osteometric Variation of the Human Spine 6 Figures

Figure 1: Anatomy of the intervertebral foramen .19

Figure 2: CT based 3-D-surface reconstructions of thoracic and lumbar spines of

selected ape cadavers .85

Figure 3: Major samples examined ..110

Figure 4: Historic age groups (time groups) represented in the whole sample ...... 110

Figure 5: Major time groups represented in the whole sample ...... 111

Figure 6: Age groups distribution in the whole sample ...... 111

Figure 7: Map of Central-'Western Europe with sample origins ...... 118

Figure 8: Lateral views of measurement definitions of the intervertebral foramen ....I24

Figure 9: Cranial views of osteometric measurement definitions of the intervertebral

foramen 126

Figure 10: Technical error of measurement for selected spinal dimensions by

experienced and inexperienced observers ...... 133

Figure 11: Sagittal diameter of Ll vertebral body in females showing mostly a normal

distribution 140

Figure 12: Transverse process width of L5 in females showing a non-normal

distribution 140

Figure 13: Transverse spinal canal diameters at Ll in males showing mostly a normal

distribution ...... 141

Figure 14: Spinous process lengths at C3 in males showing a non-norrnal distribution 141

Figure 15: Variables by vertebral levels with mean for males and females and maximum

one standard deviation range ...... 145

F. J, Rühli - Osteometric Varialion of the Human Spine 7 Figure 16: Selected variables with signifrcant change with known individual age rn

modern samples .... 155

Figure 17: Selected variables with signifrcant microevolutionary trends in males ....159

Figure 18: Selected variables with signifrcant microevolutionary trends in females ....161

Figure 19: Significant and borderline alterations of mean values between Neolithic /

Bronze Age and modern samples ... r70

Figure 20 Signifrcant and borderline alterations of standard deviations values between

Neolithic lBronze Age and modern samples ...... 174

Figure 21: Principal components I and 2 of males and females in modern samples ....I77

Figure22: Mean humerus length of various historic and modern samples ...... 246

Figure 23: Mean femur length of various historic and modern samples .....247

Figwe24: Estimated male stature of various historic and modern samples ...... 248

Figure 25: Estimated female stature of various historic and modem samples ...... 249

Figne26: Means of measured long bones by time periods in the present study ...... 250

Figwe2T: Secular trends of male and female spinal dimensions ...... 262

Figure 28: Microevolutionary trends of selected vertebral measurements ...... 263

Figure 29: Selected variables in males with signifrcant alterations with time before

present and independent of maximum femur length .264

Figure 30: Selected variables in females with significant alterations with time before

present and independent of maximum femur length ...265

F. J. Rühli - Osteometric Variation of the Human Spine 8 Abstract

For most parts of the human body, the morphometry and its variation with regard to microevolutionary and secular trends, sexual dimorphism and individual aging are well known. Surprisingly, studies focusing on the vertebral column have so far primarily used either a macroevolutionary or a clinical focus. The aim of this study is to address the osteometry and variation of the human spine from a special perspective, possible microevolutionary alterations.

A total of 348 human skeletons, dating from 28,000 B.C. to the mid 20* century

4.D., from 24 sites mostly in Switzeriand and Southem Germany, and without macroscopic pathology, were measured with a caliper by a single observer. These measurements atvertebral levels cervical 3 and 7, thoracic 1,6 and 10, andlumbar I and

5 were taken: ventral and dorsal vertebral body height, sagittal and transverse vertebral

body and spinal canal diameters, spinous and transverse process length, pedicle height

and intervertebral foramen widths; as well as the diameters of the foramen magnum,

humerus and femur length and circumference, femur head breadth and bi-iliac widths.

With the exception of most of the bony outlines of the neural pathways, males

show larger osteometric dimensions than females. No side difference of bilaterally

measured variables was found. Variables of neighbouring vertebrae correlate to a higher

extent than more distantly located variables; similar measurements at different vertebral

levels correlate generally better than non-related measurements. With greater individual

age, especially in males, the diameters of the vertebral body and pedicle height increase.

A positive microevolutionary trend, with both increasing mean values and standard

deviations, could be found; this trend was independent of stature for selected measures.

F. J. Rühti - Osteometric Variation of the Human Spine 9 The samples show a microevolutionary increase in most of the spinal variables.

Since both, mean values and standard deviations, increased, one may explain this higher intra-group variabilþ to be a result of relaxed natural selection. Various environmental or genetic factors could explain the short-term alteration of the spinal osteometry'

Furthermore, the relative smaller size and decrease with age of the bony outline of the neural pathways in males, could explain their higher vulnerability to modem lower back pathologies.

F, J, Rühli - Osteometric Variation of the Human Spine 10

Acknowledgments

"May those deceased individuals, whose sleep was disturbed

driven by scientific curiosity, rest forever peacefully!"

I would iike to express my deepest gratitude to my beloved parents for their

unrestricted support. This work is dedicated to them'

Furthermore, I would like to cordially thank

- prof M. Henneberg, Department of Anatomical Sciences, The University of Adelaide,

for being a superb supervisor, acting as a scientihc role model and, together with his

wife, Dr R. Henneberg, facilitating and enriching my stay in Adelaide

- prof M. Schultz, Department of Anatomy, Georg-August-University Göttingen, for

being a helpful co-supervisor and providing generous access to skeletal samples

- prof B. Rüttimann, Medizinhistorisches Institut und Museum, IJniversität Zürich, for

support by employing me at his institution during part of the data collection

- prof H. de Lumley and Dr E. Herrscher, lnstitut de Paléontologie Humaine, Paris; Prof

M. Teschler-Nicola and Dr K. Wiltschke-schrotta, Abteilung Archäologische Biologie 'Wien; und Anthropologie, Naturhistorisches Museum, Dr B. Kaufmann,

Anthropologisches Forschungsinstitut, Aesch; Prof A. Langaney and Dr P. Mennecier,

Laboratoire d'Anthropologie biologique, Musée de l'Homme, Paris; Prof A. Gallay

and Dr C. Kramar, Département d'Anthropologie, Université de Genève; Prof G.

Grupe, Institut für Anthropologie und Humangenetik, Universität München and Dr P.

Schröter, Staatssammlung für Anthropologie und Paläoanatomie, München;

Kantonsmuseum Basel-Land, Liestal; Prof R. D. Martin (now at Academic Affairs,

12 F. J. Rühti - Osteometric Variation of the Human Spine Institut und Fieid Museum, Chicago) and Ms E. Langenegger, Anthropologisches

Basel- Museum, universität zùnch; Ms L. Meyer, Archäologische Bodenforschung

and Dr G' Hotz, Abteilung Stadt, Basel and dipl. nat. M. Lörcher, Dr F. Wiedenmeyil

Anthropologie, Naturhistorisches Museum, Basel; Dr J. Tauber, Kantonsarchäologie, phil. F.-X' chauvière' Kantonsmuseum Basel-Land, Liestal; Prof D. Ramseyer and lic' for their lnstitut de Préhistoire, université de Neuchâtel and Latènium, Hauterive,

staff for technical help hospitable help and providing access to the collections and their 'Washington St' Louis; - Prof E. Trinkaus, Department of Anthropology, University, New Orleans; Prof T. w. Holliday, Department of Anthopology, Tulane university, universþ, Athens Prof N. Tatarek, Department of sociology and Anthropology, ohio

and Evolution, John Hopkins and Mr B. M. Auerbach, centre for Functional Anatomy

university school of Medicine, Baltimore for providing me with comparative

measures, additional collection data or literature references

sciences, The - T. Morgenstern and other members of the Department of Anatomical

University of Adelaide, for technical support

Research - the university of Adelaide for awarding me an "International Postgraduate

Scholarship" to hnancially allow this study

,,Dr. the CT scans of the - the staff of Jones and partners", Adelaide, for performing

gteat aPes and Industrial - the l2,h-year-students involved in the "Commonwealth Scientifrc for their participation in Research organisation,' Student Research Scheme; Adelaide,

the project on reliability of osteometric measurements

and my siblings Dr M' B'Mettler- - and last but not least my girlfriend Ms c. F. Gassiot

Rühli and Dr M. E. Rühli for their permanent morale encouragement!

13 F. J. Rühti - Osteometric Variation of the Human Spine Introduction

Anatomical aspects of the human spine

The spine is a crucial and individually different part of the human axial skeleton.

As Bohart (1929, p. 698) mentioned: "...an individual's spinal column ,s cts characteristic of that individual as his face...". Similarly, Ravenel (1877) already mentioned the high degree of inter-individual variability of the human spinal dimensions.

The anatomical structure of the human spine has been studied on a macroscopic and microscopic level for many centuries. To be able to distinguish between the occurrence of an abnormality and an anatomical variation within the human vertebral column, one has essentially to conduct a precise assessment of the normal structure and its size. This assessment can be done by various approaches, either by using animal models (Iwamoto et al., 1995), in clinical studies involving asymptomatic and / or

symptomatic patients (Horner, 1854; Blumensaat and Clasing, 1932; Junghanns, 1933;

Elsberg and Dyke, 1934; Wolf et al., 1956; Epstein et al., 1962; Burrows, 1963;

Hunrthal, 1968;Katz et al.,1975; Ramani, 1976; Portet et al.,1978a; Porter et al.,1978b;

MacGibbon and Farfan, 1979;Larsen and Smith, 1980a; Larsen and Smith, 1980b; Porter

et a\.,1980; Stockdale and Finlay, 1980; Ullrich et al., 1980; ogino et al., 1983; \üeisz

and Lee, 1983; Drinkall et a1.,1984; Kikuchi et al., 1984; Macdonald et al., 1984; Nissan

and Gilad, 1984; Bolender et al.,1985: van Schaik et a\.,1985; Gilad and Nissan, 1986;

Nissan and Gilad, 1986; Gallagher et al., 1988; Hedlund and Gallagher, 1988; Minne e/

al., 1988; Davies et al., 1989; Black et al., 1991; Hermam et al., 1993; Frobin et al.,

1997; Humphreys et a\.,1998; Wildermuth et a\.,1998; Schmid et a\.,1999; Harrington

et a1.,2001), by using cadaver material (Horner, 1854; Ravettel, 1877; Jacobi, 1927;

F. J. Rühli - Osteometric Variation of the Human Spine 74 Larmon, 1944; Magnuson, 1944; Dommisse, 1974; Dommisse, 1975; Veleanu, 1975;

Crock, 1981; Hasue et al.,1983; Bose and Balasubramaniam, 1984; Kikuchi et al.,1984;

Rauschning, 1987: Stephens et al., 1991; Yoo et al., 1992; Hasegawa et al', 1995;

Ebraheim et a1.,1996; Nowicl

Veleanu, 1975; Eisenstein, 1977; Kikuchi et al., 1977; Eisenstein, 1980; Postacchini e/ al., 1983; Berry et a\.,1987; Scoles et a1.,1988; Lee et a1.,1995; Ebraheim et al.,1996;

Cinotti et aL.,2002).

The measurement of the human vertebral column has been so far def,rned for radiological (Elsberg and Dyke, 1934; Wolf et a1.,1956; Bunows, 1963; Hurxthal, 1968;

Jones and Thomson, 1968; Yttal et al., 1983; Nissan and Gilad, 1984; Bolender et al.,

1985; van Schaik et a\.,1985; Gilad andNissan, 1986; Nissan and Gilad, L986;Kraget al., 1988; Marchesi et a\.,1988; Olsewski et al., 1990; Stephens et ql., l99l;Yaccaro et al., 1995; Kothe et a1.,1996; Karaikovic et al., 1997; Schmid et a1.,1999;Harnngton et al., 2001; Kandziora et a1.,2001) or osteometric studies (Horner, 1854; Aeby, t879;

Anderson, 1883; Rosenberg, 1899; Wetzel, 1910; Hasebe, 1913; Thomson, 1913; Cyriax,

1920; Stefko, 1926; Jacobi, 1927; I|rlafün, 1928; Huizinga et al., 1952; Veleanu, 1972;

Veleanu, 1975; Saillant, 1976; Kikuchi et al., 1977;Put2,1981; Postacchini et al.,1983;

Larsen, 1985; Cottenll et a\.,1986; Berry et a\.,1987; Marchesi et al.,1988; Scoles et al.,

1988; Olsewski e/ al., 1990; Gepstein et al., 1991; Panjabi et al., l99la; Panjabi et al.,

I99lb Panjabi et al., 1992;Ho,l et a1.,1993; Shapîo, 1993; Shapiro, 1995; Tominaga et al., 1995; Vaccaro et a\.,1995; Xu et al., 1995; Karaikovic et al., 1997;Kandziora et al.,

2001; Cinotti et a1.,2002).

F. J. Rühti - Osteometric Variation of the Human Spine 15 The normal human spine consists, besides the sacrum and the coccyx, of 24 vertebrae: seven cervical (abbreviated: Cl - C7), twelve thoracic (Thl - Th12) and five lumbar ones (Ll - L5). The vertebrae enclose the spinal cord, which usually ends between

Ll and L2 (McCotter, 1916i.

'Whereas the major function of the vertebral bodies is to carry the body weight and serve as an axis for body mechanics, with the intervertebral discs acting as buffers, the main purpose of the vertebral arch, the pedicles and the laminae, is to protect the spinal cord and to link with the transverse and spinous processes, which serve as the attachment points of various supportive back muscles. The spinous processes also limit, together with the ligamentaflava, extension movements at least of the human thoracic spine (White and

Hirsch, l97l).

The spinal cord consists of the grey matter, the nerve cell bodies, and the white matter, containing the nerve fibres. Nerve roots exit from the spinal cord on each vertebral level and provide sensory and motor innervation to the periphery. The white matter includes the dorsal columns, linked with sensory abilities, and the latero-ventral columns, which represent the motor innervation.

The neural canal contains the spinal cord and its nerve roots, the cerebrospinal fluid, the dural sac, extradural fat, ligaments and, just behind the vertebral bodies, a venous plexus. Furthermore, a menigeal recurrent nerve providing nociception to the ligaments, the spine, the dura and the vertebrae can be found in this area. The dural sac extends further caudally and ends mostly on sacral (S) level I or 2 (Salamon et al.,1966).

The particular spinal neural situation was reviewed earlier (Rydevik et al., 1984;

Group and Stanton-Hicks, 1991) as well as the spinal ontogeny and adult anatomy

(Donaldson and Davis, 1903) and its aging related adaptation (Bailey, 1953). Larsen

F. J. Rühti - Osteometric Variation of the Human Spine 1'6 (1985) already discussed widely the specific anatomical interaction of the lumbar spinal nerves and the posterior surface of the vertebrae. He even mentions the fact that the postero-lateral vertebral body parts develop from the same ossification centre, as do the spinal neural arches.

The spinal cord is surrounded among others by its meninges and peridural flat.

Between the spinal cord and the osseous and ligamentous borders of the spinal canal, a free space, so called "spinal canal reserve capacity" (Weisz and Lee, 1983), is located, which allows the spinal cord to move freely and independently from body movements.

The anatomy of the intervertebral foramen in relation to its surrounding osseous and soft tissue structures has already been widely addressed (Swanberg, 1915; Larmon,

1944; Magnuson, 1944; Epsteín et al., 19641' Veleanu, 1975; Crock, 1981; Kirkaldy-

Willis et al., 1982; Hasue et al., 1983; Vital et al., 1983; Bose and Balasubramaniam,

1984; Kikuchi et al., 1984; Vanderlinden, 1984; Rauschning, 1987; Hoyland et al., 1989;

Mayoux-Benhamou et a\.,1989; Stephens et al.,I99l; Hasegawa et aL.,1995; Ebraheim et al.,1996). Rauschning (1987) describes the outline of the lumbar root canal as being of an inverted teardrop form with an oval shaped intervertebral foramen at its caudal end.

Hasue et al. (1983) characterize the normal form of the lumbar intervertebral foramen as being oval or almost triangular at least in the cadaveric spine. Bose and Balasubramaniam

(1984) call the intervertebral foramen the "external ring" of the nerve root canal with oval size for the two lowest lumbar levels and more circular shape for Sl.Lee et a/. (1988) divide the lateral section of the lumbar spinal canal into three major parts: The entrance zone containing the nerye root and the dura mater; the mid-zone which consists of the dorsal root ganglion, which is usually located in the supero-lateral area and often plays a 'Weinstein, significant role in lower back pain symptoms (Vanderlinden, 1984; 1986;

F. J. Rühti - Osleometric Variation of the Human Spine 77 Hasue et a1.,1989), and the ventral motor nerve root surrounded by fibrous extensions of the dura mater and, finally, the exit zone with the peripheral nerve and its perineurium cover. Vital et al. (1983) divide the lumbar radicular canal also into three morphologically different sections, which are the retrodiscal space, the parapedicular space or lateral recess and, the intervertebral foramen. The major factors affecting the intervertebral foramen size are e.g., degenerative changes of the bony borders, increased spinal mobility, disc degeneration, subluxation of the facet joints or bulging of the ligamentum flavum. The intervertebral foramen, the exit zone according to the categonzation by Lee et a/. (1988), contains beside the spinal nerye, which is mostly located in its inferior section, only fat and blood vessels in the upper section of the foramen (Swanberg, 1 9 I 5).

Surprisingly, most of the intervertebral foramen seems to be filled out by fat tissue, which acts, due to its semi-liquid consistence in living people, as a natural buffer for any physical stress operating on this anatomical region and, in particular, the exiting nerve roots (Swanberg, 1915). According to Swanberg (1915), who examined the microscopic structure of the intervertebral foramen, the neural tissues in the intervertebral foramen also show a lack of major lymphatic vessels. Hoyland et al. (1989) investigated the normal and clinically abnormal microscopic structure of the intervertebral foramen and describe its main content similar to previous reports. In a normal intervertebral foramen, the fibrous tissue occupies less than 28o/o,the neural tissue less than 35o/o and a lot of vessels of diverse sizes were found (Hoyland et a1.,1989). The foramen, according to them, forms an outline of an upside-down pear. Some of the earlier described outlines of the intervertebral foramen could be found in Figure 1.

F. J. Rühli - Osteometric Variation of the Human Spine 18 A B

lnverìÈd pear Tear dtop

LF F va

cap. D

VB P

ovãl Auri€ulâl

c

Figure i Anatomy of the intervertebral foramen (hgures unchanged or slightiy modifred from original references)

root of the A) Intervertebral foramen of the cervical spine (Veleanu, 197 5); 1: spinal ganglion, 2: anterior nerve, 5: vertebral spinal nerve, 3: anterior ramus of the spinal nerve, 4: posterior ramus of the spinal 9: spinal artery,6: vertebral periarterial venous sinus, 7: cervical epidural venous sinus, 8: dura mater, cord, l0: unciform process, 11: upper articularprocess B) Bony and cartilage outline of the intervertebral foramen (Swanberg, 1915); A: inferior articular E: head process, B: root of superior vertebral arch, C: verlebral body, D: intervertebral fibro-cartilage' of rib, F: root of inferior vertebral arch, G: superior articular process arch, V[l: vertebral C) Intervertebral foramen shape (Stephens et a1.,1991); P: pedicle of the vertebral bodies above and below, D: intervertebral disc, LF: ligamentum flavum, CAP: capsule of

zygoapophysial joint, IAF / SAF: inferior and superior articular facets

19 F. J. Rühli - Osteometric Variation of the Human Spine A precise knowledge of the anatomical peculiarities of the human vertebral column is crucial to understand not only its special purpose and to help to evaluate its specific evolutionary background, but also to understand some particular clinical problems.

The clinical significance of the particular anatomy of the lower human spinal column has already been discussed in detail by Magnuson (1944). Crock (1981) described the anatomy and its linked pathology of the lumbar spinal nerve root canal, which was also reviewed by Rydevik et al. (1,984). Crock (1981) emphasizes the importance of the

"spinal nerve root canal concept" especially for the lowest lumbar region, rather than the use of the term "lateral recess" which, according to him, is just true and useful for a minor part of the spinal nerve pathway. Furthermore, Bose and Balasubramaniam (1984) discuss the particular anatomy of the lumbar nerve roots canals. They introduce the term

"external ring" for the exit at the intervertebral foramen. They also provide, besides detailed anatomical descriptions, measurements of the nerve root canal lengths. Veleanu

(1912; 1975) addressed the particular anatomy of the cervical nerve root grove and the unco-transversal region. For the cervical nerye groove, he differentiates two parts, the initial radicular part and the terminal groove of the anterior spinal nerve ramus.

The particular situation of the lumbo-sacral dural sac and the linked nerve roots has been discussed widely by Salamon et al. (1966) and the anatomy of the lumbar nerve root canals has been highlighted by Bose and Balasubramaniam (1984). Nerve roots in the spinal column have no perineurium, weakening them in strength in comparison to other peripheral nerves (Sunderland and Bradley, 1961) and implying a higher susceptibility to compression. Nevertheless, Hasue et al. (1983) and Kikuchi et al. (1984) mention an epiradicular membranous layer around the exiting nerve root and Hoyland e/

F. J. Rühl¡ - Osteometric Variation of the Human Spine 20 al. (1989) describe the dura mater covering the nerve roots at the entry of the

intervertebral foramen. Nerve roots hll out approximately 20-50% of the intervertebral

foramen dimension and lie anterior to the dorsal root ganglion, which is also a part of the

lateral intervertebral foramer and consists of the cell bodies of the sensory neurons

(Swanberg, 1915; Bornstein and Peterson, 1966; Panjabi et a\.,1983; Vital et al., 1983;

Bose and Balasubramaniam, 1984; Vanderlinden, 1984; Hoyland et al.,1989; Hasegawa

et al., 1995). Magnuson (1944) states that the nerve ganglion is just slightly smaller than

the intervertebral foramen, both apparently measuring 7 mm in average atL4 and L5. The

exiting spinal nerve roots pass just below the pedicle of the upper vertebral level, in the

upper part of the intervertebral foramen (Kirkaldy-Willis et al.,1982; Rauschning,l9ST).

Larsen (1985) highlights the fact that the lumbar nerye roots are even more flexible than

the more cranial ones due to their longer intraspinal segments, which may have an impact

on the infero-lateral posterior vertebral surface.

All these facts have clinical relevance, as discussed later. This is particularly true

for example for the interaction between the osseous- and non-osseous parts of the

intervertebral foramen and its corresponding nerve roots, which according to Mayoux-

Benhamou et al. (1989) are key factors in such clinical situations.

Another osteometric landmark of the neural spinal canal is the foramen magnum,

located at the skull base. Schaefer (1999) found e.g., that the distance of the foramen

magnum from the bi-carotid chord could be used to differentiate human from non-human

e.g., chimpanzee, crania. Nakashima (1986) compared types of size of the foramen

magnum in male middle Kyushuites with that of male Germans and postulates a possible

change of type during individual growth. Nakashima (1986) found a variance in length

but not in breadth of the foramen magnum among these groups. Martin (1928) stated that

F. J. Rühti - Osteometric Variation of the Human Spine 27 there is a high individual variability in foramen magnum dimensions. The main diameters of the foramen magnum have already proven their possible predictor qualities for body mass, not only in humans but also in other hominids (Aiello and Wood, 1994).

Furthermore, foramen magnum size does not correlate well with spinal cord size in primates but with body weight (Maclarnon,1996a).

The osteometry of the major spinal regions has been widely covered in the studies by Panjabi et al. (I99la; l99lb; 1992). Based on their morphometric research they

declare the following major spinal zones to be of transitional nature: C2 - C3, C6 I C7-

Thl, Thl - Th4, Th10 - Thl2,Ll -Lj,L3 -L5. Furthermore,Putz (1981) investigated

and reviewed the major aspect of the spinal anatomy, its ontogenetic development and the

functional anatomy with special focus on the spinal joints.

The ontogeny of the human spine has already been addressed by Aeby (1879). He

found e.g., that adults have a relatively longer lumbar spine but shorter cervical spine,

with the thoracic spine being relatively similar to its size in childhood. According to

Aeby (1879), the spinal canal dimensions change remarkably by becoming relatively

smaller in adulthood. Additionally, the adult spine is slimmer in the transverse plane

(Aeby, 1879). Also Donaldson and Davis (1903) as well as Lassek and Rasmussen (1938)

reported the ontogenetic aspects and the adult anatomy of the spinal cord. According to

Donaldson and Davis (1903), the majority of the spinal cord area increase occurs after the

age of five years, with a relative higher increase of the white matter and more prominent

change in the thoracic region. Lassek and Rasmussen (1938) describe a relatively bigger

increase of the white matter and of the thoracic spinal cord as well as a relative shrinking

of the spinal cord length from the newborn to to adulthood; with the average spinal canal

length being 410 mm. Donaldson and Davis (1903) found an average length of the spinal

F. J. Rühli - Osteometric Variation of the Human Spine 22 cord in adults of 441 mm, with an indication for a conelation between the length of the

vertebral column and the osseous spine.

Furthermore, Donaldson and Davis (1903) found differences in adults between

spinal cord cross-section areas and volumes of the grey and white matter at various spinal

levels. Whereas the maximum spinal cord area locates in a mixed sex sample at level C6,

followed by the values on level L3 and L5, the ratio between white and grey matter varies

depending on vertebral level, with the grey matter usually being approximately 20Yo of

the white one (Donaldson and Davis; 1903). Lassek and Rasmussen (1938) found an

average ratio of approximately 18%. The highest volume of grey matter is found,

according to Donaldson and Davis (1903), at the level C6, whereas the highest value of

grey matter area can be found at level L5. Donaldson and Davis (1903) further explored

the relation between grey matter and spinal nerves at various vertebral levels. They found

that only for the cervical and sacral region there is a correspondence between the two,

whereas for the thoracic and lumbar section there is not. In the latter two, the grey matter

volume is bigger than expected, explained by them as a reaction of vertebral elongation

during growth rather than increased neural complexity at these levels. McCotter (1916) reports an average spinal cord length for White males of 448 mm and for'White females

of 418 Írn, as well as 434 mm in Black cadavers with unlisted sex. Furthermore, Ravenel

(1877) reports lengths of the total vertebral column for fresh male and female cadavers.

He highlights the high degree of inter-individual variability of spinal osteometric measurements, which reaches in some dimensions up to a third of the mean value.

Surprisingly, the juvenile spine reaches at a very early age most of its adult

dimensions (Porter and Pavitt, 1987). However, the juvenile spinal canal still further changes its shape by maturing (Porter and Pavitt, l98l). Wolf e/ al. (1956) state that the

F. J. Rühli - Osteometric Variation of the Human Spine 23 lower clinically critical limits of the sagittal spinal canal dimensions are attained at the age of four to five years, whereas the adult size, according to them, may be reached at an average age of 12 years. Clark et al. (1985) state that approximately 90o/o of the adult vertebral canal size is completed by late infancy, which makes the spine more vulnerable to prenatal growth disruptions than other parts of the human body.

Various reports already addressed the relationship between spinal cord size and brain size (Marshall, 1892; Latimer, 1950; Maclarnon, 1996b). Latimer (1950) reports

for Guinea pigs correlations between spinal cord weight and total brain weight as well as

the weight of various brain parts, between spinal cord weight and length and between

spinal cord weight and both body weight and length. Latimer (1950) found weaker

correlations for the spinal cord length than weight e.g., in relation to body weight.

Maclarnon (1996b) describes also a correlation between brain weight and spinal cord

weight for primates. Marshall (1892) calculated the spinal cord to be 2.1 % of the brain

weight in humans. Maclarnon (1996b) lists, based on own studies and summarized from

earlier published data, an average brain weight of an adult 60 kg human tobe 1274 g and

a spinal cord weight of 29.7 g. These values are from unpublished data sources by Martin

and Maclarnon and differ from established averages of approximately 59 kg and brain

weight of about 1350 g (Pakkenberg and Voigt, 1964; Beals et al., 1984; Henneberg,

1990), and, therefore, some caution is necessary for these data. The overall spinal cord

length was reported in this study to be 448 mm in males and 413 mm in females

(Maclarnon, 1996b). Nevertheless, Elliot (1945) found no evidence for a correlation

between individual stature, and weight, as well as sex or age and spinal cord dimensions

in humans.

F. J. Rühli - Osteometric Variation of the Human Spine 24 The size of the vertebrae and its use for estimation of body size is still doubtful

(Martin and Saller, 1957; Tibbetts, 1981). Tibbetts (1981) found that the coefficient of correlation mostly increases the larger the vertebral numbers included in individual stature estimation are. Gozdziewski et at. (1976) found a clear correlation between thoraco-lumbar spine length and individual height in a sample of living individuals.

Karaikovic et al. (1997) describe a dependence of pedicle diameters on individual body height of 30o/o up to 70%o.In a modern Polish medical student sample individual height was signifrcantly correlated with the length of the thoraco-lumbar spine (Gozdziewski e/ at., 1976). Also, Minne et al. (1988) mention a clear correlation between individual stature and vertebral dimensions. In a radiographic study on living women, Gallagher et a/. (1988) found a correlation between thoraco-lumbar vertebral anterior and posterior height and individual height or weight. Amonoo (1985) found a change in the mid-

sagittal neural canal diameters in relation to alterations of the sagittal diameter of the vertebral body. For both sexes, he mentions for such a ratio a value of 0.5 onL2 - L5 and

0.6 on L1 respectively.

On the other hand, in the osteometric study published by Berry et al. (1987), the

combined vertebral body heights did not show a correlation with the recorded individual

height of the deceased person. Piera et ø/. (1988) found no correlation between pedicle

transverse diameter and its equivalent dimension of the neural canal. Katz et al. (1975)

found no influence of stature on human cervical vertebra morphometry. Contrary, the

"ponderal index", which is body height divided by'{ body weight, as well as the head

weight correlated at least with some of the cervical spine measurements.

The influence of body size and weight on spinal morphometry, especially the size

of the neural canal diameters has akeady been addressed (Legg and Gibbs, 1984;Porter et

F. J. Rühti - Osteometric Variation of the Human Spine 25 gton et al. 2001) Porter et al' (1987) al., 1987 ; Sanders, 1 99 I ; Maclarnon, 1995; Hanin , ' found that patients with a narro\il sagittal spinal canal diameter were 22o/o heavier than their counterparts with a wide one. Body weight is proposed to be the best variable reflecting body size (Jungers, 19S4). Harrington (2001) did not find a correlation between individual stature, weight and body mass index for the occulrence of a disc hemiation in the lower lumbar region. On the other hand, Heliövaara (1987) describes a link between stature and in particular moderate increase in body mass index and, only in males, the hospitalisations due to herniated lumbar discs. Legg and Gibbs (1984) found no clear correlation between individual stature or body weight and lumbar spinal canal

dimensions. Furthermore, Murrie et al. (2003) found a more prominent lumbar lordosis in

individuals with a higher body mass. Furthermore, body weight seems not to be related

with the dorsal root / ventral root ratio in the spinal canal (Corbin and Gardner, 1937).

Nevertheless, body weight in mammals is correlated with the size and number of spinal

nerve root fibres (Dunn, l9l2).

In an archaeological sample, Hibbert et al. (1981b) describes a correlation for

individual long bone sizes and transverse spinal canal diameter, but not for its sagittal

counterpart. Additionally, Porter and Pavitt (1987) v/ere unsure about the direct influence

of small transverse juvenile spinal canal diameters, the known stress marker dental

hypoplasia and individual stature. Jankauskas (1994) reports a correlation of less than 0.4

between individual stature and longitudinal spinal measurements. Furthermore for a

female sample, Ross e/ at. (1991) could not detect a signihcant change in vertebral

morphology such as anterior / posterior vertebral body height ratio with individual

stature. McCotter (1916) did just find a tendency but no clear correlation between spinal

cord length and individual stature as measured by height or vertebral column length. In a

26 F. J. Rühli - Osteomelric Variation of the Human Spine clinical study by Harrington et at. (2001), individual height, weight or body mass index did not have an influence on the occurrence of disc herniation. Nevertheless, there was a

a just very correlation between vertebral body diameters and individuai body weight, but weak one with pelvic breadth.

sexual dimorphism and age-related adaptations of the human spine

It is well known that sex influences the spinal morphology, since e.g., already

Dwight (1894) reported longer relative lumbar regions in women' Martin (1928)

than its male mentions, that the female ventral vertebral body height is usually smaller

upper thoracic counterpart, with an especially prominent difference for the cervical and of region. MacGibbon and Farfan (1979) found no influence of sex on the occurrence

the transverse transitional lumbar vertebra and rudimentary ribs. However, they found

This would process atL5, in relation to the reference one at L3, to be longer in females' atL5 I predispose females to have more likely degenerative changes atL4 lL5 and males

spinal canal S1 (MacGibbon and Farfan, l97g). The overall morphology of the lumbar

(1985) does not show any sex dependent variation (Piera et al',1988). Amonoo-Kuofi

and generally more describes in a study of vertebral columns from Nigeria a nalrower

(1955) reports to have variant sagittal diameter of the neural canal for females. Francis relative found only absolute smaller values for female spines, but without any appafent

Furthermore,}i4ltta et alterations of the vertebral dimensions in relation to male samples.

to sex, mostly al. (2002) found only non-signif,rcant differences in pedicle size in relation

(1993) orXu et al' similar to the resuits presented by Ebraheim et al. (1997),Hou et al. in (1995). In addition, Karaikovic et al. (Igg7) did not find sex-dependent differences

of the pedicle dimensions, once the influence of different body height was taken out

27 F. J. Rühli - Osteometríc Variation of the Human Spine calculations. Nevertheless, for the pedicle height, Olsewski et al. (1990) found a signif,rcant sex difference, with women having smaller dimensions than males, for most lumbar levels. Burrows (1963) could not find a significant difference in sagittal spinal canal diameter between sexes, he describes a difference of usually I mm or less. No significant differences have been shown in foraminal dimensions in relation to sex

(Ebraheim et al., 1996). Also Hinck et al. (1966), in their study of the interpediculate distance as observed on roentgenograms, stress the only minor influence of sex on at least a selection of spinal dimensions. In a biomechanical study, Nachemson et al. (1979) found no correlation between age or degenerative lower back pathologies and altered mechanical behaviour of the lumbar motion segments. Furthermore, they only describe a

slightly higher flexibility of female motion segments to bending or compression forces.

Sex differences were also found by Tatarek (2001) with larger neural lumbar canals in

males than in females. This is in general consistent with the findings reported by Lee et

al. (1995) for the mid-sagittal diameter but not for the interpedicular diameter.Píera et al.

(1988) found in an X-ray based study, that there is a link between sex and interpeduncular

distance of lumbar Ll - L4. Katz et al. (1975) found in another X-ray based study on

recent volunteers, that males have signihcantly larger cervical vertebrae height and

sagittal width than females. These findings by Katz et al. (1975) -ay be caused by the

fact that for the two sexes individuals of the same percentile and not absolute stature were

chosen and, therefore, males were bigger on average. Van Schaik et al. (1985) found

smaller osteometric length values in females, but no differences in vertebral angles or

ratios. Horwitz (1939) also reported, at least for all measurements but not for the indices,

a highly significant sexual dimorphism with a general tendency of kyphosis in males in

thoracic spine. Minne et al. (1988) describe only a non-significant sexual difference in

F. J. Rühti - Osteometric Variation of the Human Spine 28 spinal morphology; such as the higher lumbar increase in vertebral body height in females. Females also show a signihcantly more prominent lumbar lordosis (Murrie e/ al., 2003). ln an osteometric study on the lumbar spinal canal significant sexual dimorphism was found for the ratios of the spinal canal dimension to vertebral body

(Kikuchi et al.,1977). Men have significantly larger lower lumbar vertebral endplates but the shape ratio of them seems not to differ between sexes (Harrington et a1.,2001). In a sample of asymptomatic Polish medical students Gozdziewski et al. (1976) found a larger thoraco-lumbar spine in the male sample than in the females. Berry et al. (1987) did not separate sexes in their study on spinal morphometry. According to them, even without separating the data, sex showed a coefhcient of variation of mostly less than 10 %. In the thoracic spine, Piontek (1973) detected that females show a stronger increase caudally in main vertebral body dimensions, such as sagittal and transverse diameter. The single exception was vertebral body height (Piontek, 1973). Piontek (1913) describes also in the thoracic spine a higher enlargement of the sagittal dimension caudally than for the transverse ones, whereas for the lumbar spine this trend seems to be opposite. In the Early

Medieval samples presented by Piontek (1973) the relative increase of vertebral body

height was higher for males than females and the relative increase of the sagittal

dimension was bigger for females than males.

The earlier reports on the influence of aging on spinal morphometry are equivocal.

Age-related alterations of the spinal morphometry (Jacobi, L927;Hurxthal, 1968; Trotter

and Hixon,1.974; Ericksen, 1976; Hansson and Roos, 1980; Portet et al., 1980; Weisz

and Lee, 1983; Galtagher et a\.,1988;Piera et al., 1988; Jankauskas,1992; Hermann e/

al., 1993; Edmondston et al., 1994; Jankauskas, 1994;Diacinti et al., 1995; Jason and

Taylor, 1995; Lee et al., 1995; Humphreys e/ al., 1998; Tatarek, 2001), changes in

F. J. Rühli - Osteometric Variation of the Human Spine 29 relative spinal region length (Schultz, 1961; Jason and Taylor, 1995) and vertebral bone

mineral content (Hansson and Roos, 1980) have been reported so far. Furthermore, aging

results in a general decrease of skeletal weight, with the male bones being significantly

heavier than the female ones (Trotter and Hixon, 1974). In a radiographic study on

asymptomatic females, Gallagher et al. (1988) found no correlation of the anterior

vertebral body height and individual age, whereas the posterior height was negatively

(1989), correlated. Surprisingly, in a similar study conducted by Davies et al. no change

in either anterior or posterior vertebral body height was reported for asymptomatic

females as well as women suffering from osteoporosis within time periods of at least 10

years span shortly before menopause. contrary, Black et al. (1991) conclude that no just morphometric changes occur depending on age, while Hermann et al. (1993) found a

very weak interference. Hermann et at. (1993) even argue that the described aging effect

could be due to secular increases in individual stature within the cohort. However, no age

related changes in lumbar spinal canal dimensions were found in an osteometric study by

Kikuchi et al. (1977). Also the lumbar lordosis seems not to change with age (Murrie e/ just al., 2003). Edmonston et al. (1994), in cadaver spines of elders, found a weak

correlation of vertebral body height ratios and bone density. Bone density is

representative of bone remodelling, which in the elder spine can be present in form of

wedging and increased thoracic kyphosis. Píeru et ø/. (1938) describe the absence of any

relation between general lumbar spinal morphology and age. Nevertheless, the lumbar

interpeduncular distance in particular seems to increase with age, more prominent on the

upper than on the lower lumbar spine (Piera et a1.,1988). In the same study, a correlation

of Ll - L4 interpeduncular distance in relation to sex was found as well' By focusing on

the anatomy of the human spinal cord, Elliot (1945) found not only a high degree of inter-

30 F. J. Rühti - Osteometric Variation of the Human Spine individual variation, but also independence of individual sex, age or weight from the cord dimensions. ln addition, Bailey (1953) did not find any major atrophy of the spinal canal in the elderly. However, in a study on albino rats, Dunn (1912) reports a clear decrease of nerve tissue with older age in the cervical nerve root.

The particular aspect of spinal ontogeny was highlighted, as discussed above, by

Donaldson and Davis (1903). ln addition, Aeby (1879) emphasizes the ontogenetic impact on relative dimensions in the vertebral column. Jacobi (1927) describes in his cadaver series an increase in vertebral body heights within the young adult age group, most likely due to still ongoing vertebral growth. Furthermore, he found for most anterior and posterior vertebral body heights a decrease for the oldest group, aged 70 and above.

Similar age related changes were addressed by Hurxthal (1968) with in particular an anterior decrease in vertebral body height in the elderly and a widely seen slight wedging of the dorsal part of the vertebrae. For the spinal cord of the elderly, Bailey (1953) did not find a decrease in size nor frequent thickening of the meninges, but occasionally calcareous deposits and quite often mild arteriosclerosis. Burrows (1963) could not find a change in cervical sagittal spinal canal dimensions with age. The influence of aging and menopausal status was examined by Diacinti et al. (1995). According to them, in the

female spine there is a decrease in vertebral body height of approximately 1.5 Íìm per

year with a more prominent trend for the anterior part of the vertebra.

Biomechqnics and spinal morphologt

The unique stability and instability of the human spine was discussed among

others by Louis (1985). His proposed classic "three-column" theory of spinal stability is

in accordance with the normal ossification pattern of the spine. The first pillar is the

F. J. Rühti - Osteometric Variation of the Human Spine 31 vertebral body, whereas the second and third ones are formed by the posterior articular processes, all of them resisting the forces of gravity (Louis, 1985). Louis (1985) found an increased size of the three pillars and of the flexor and extensor trunk muscles caudally.

According to his model, the vertical axial stability is maintained by the three pillars, which are toughened by the horizontal vertebral arch. Louis (1985) attributes the spinous processes no role in maintaining spinal stability. Transverse stability of the spine is reinforced by bony and ligamentous stabilizers, varying for flexion, extension or rotation movements (Louis, 1985). Louis (1985) describes the spinal segment units as consisting of three joints, the intervertebral disc and the two zygoapophysial joints; the latter ones orientated at a different angel to the disc and supporting the weight bearing' Depending on the body position and physical load, either the disc of the two posterior joints resists compression forces with the other one resisting shearing impact (Louis, 1985).

The important role of the so called posterior elements, consisting mainly of the facet joints, parts of the laminae, the spinous processes and some ligaments, was examined by White and Hirsch (1971) by showing the biomechanical result after the ablation of these structures. Putz (1981) not only describes the main osseous aspects of the human spine but focuses especially in his study on the anatomical and functional particularities of the vertebral joints as they act in collaboration with other bony

structures, ligaments and muscles. He divides the human spine in various

"Bewegungsregionen", motion regions, which show a different active and reactive

pattern at the various positions and forces acting on them. These segments stretch,

according to him, from Cl to C3, C3 to Thl(2),Thl(2) to Th(l1)12, and Th(11)12 to the

end of the sacrum, with the thoracic part consisting of two major regions, Thl to Th8 and

Th8 to Thl2. Putz (1981) widely discusses the functional implications of the particular

F. J. Rühti - Osteometríc Yariation of the Human Spine 32 anatomy of the vertebral joints. He describes the axial pressures to be transmitted to the spine in three main axes, the vertebral body and the two vertebral joints. The importance of the zygoapophysial joints in maintaining spinal stability was also highlighted by Putz

(1981). Veleanu (1972;1975) highlights the importance of the'hnco-transverso-articular- complex" in particular for the mechanical stability of the cervical spine, with the transverse epiphysis as a blocking factor preventing mechanical instability and protection for the neural and vascular contents of the neural pathways. Schmorl and Junghans (1968) used the term "motor segment" to describe all the soft tissue linking the disc and the apophyseal j oint complex.

The impact of axial loading on the human spine with a particular focus on the posterior vertebral body and the intervertebral disc was studied by Larsen (1985). The concavity of the posterior vertebral body surface is explained in his model as caused by load induced traction forces as well as the pressure acting by the cerebrospinal fluid.

Adams et al. (1994) and Panjabi et al. (1976) already discussed the impact of axial loading on motion segments, which consists of two adjoin vertebra and their intervertebral discs. The lack of this axial impact on the spine in cadaveric studies is discussed as a weak methodological point e.9., by Fujiwara et al. (2001). Both, flexion and extension biomechanically influence in different ways the various spinal components.

The amount of physiological forces acting on the healthy vertebral column, even in simple movements only, is quite astonishing. Nachemson (1966) detected, in an experimental in vivo study in sitting positions, involved forces of at least twice the individual body weight above the selected mid-lumbar vertebral levels; such forces are ranging from approximately 1000 N up to 1800 N. The decrease of these forces to roughly half of their value in upright standing situation was explained by Nachemson

F. J. Rühli - Osteometric Variation of the Human Spine 33 (1966) as a result of smaller forces impacting from the muscles such as psoas major in this particular position. An even higher decrease was found in a physiological reclining movement. The load increases dramatically if one bends forward, especially with weight bearing hands. For such a situation forces of up to four times of the individual's body weight working on the lumbar intervertebral discs have been proven by Nachemson

(1966). Apparently, the ligamentous components of the spine are not strong load bearing forces, but work together with the rib cage as stabilizer of the spine (Nachemson, 1966)'

Nachemson (1966) emphasizes the extremely high shearing forces, which act on the dorsal part of the anulus fibrosus and may be of clinical significance as weli. In another biomechanical study, Nachemson et al. (1979) found no clear influence of age or sex on physical performance of lumbar motion segments. Only females seem to have segments that are slightly more flexible in response to bending and compression forces.

Furthermore, Veleanu (1972;1975) highlights in his study of macerated and cadaveric cervical spines, the importance of the tratrsverse process within the '1mco- transversoarticular" complex in limiting possibly pathologic movements. Adams et al'

(Ig94) examined the influence of flexion and extension on the various load-bearing spinal structures. Surprisingly, Adams et at. (1994) conclude that a mild flexion is the best compromise for a spinal position in weight bearing.

Osteometricfindings of earlier spinal studies

Spinal morphometry differs essentially for each vertebral level (Black et al.,l99l;

Hermann et a\.,1993) and is reported remarkably different in various studies'

The ventral vertebral body height, according to Lanier (1939), increases from C3

caudally, with the exception of C5 and C6 that have the smallest values. Hermann et al.

F. J. Rähli - Osteometric Variation of the Human Spine u (1993) report a consistent increase for both sexes from Th4 caudally. Anderson (1883) describes a decrease caudally in anterior vertebral body height for most of the cervical spine with an increase of its size caudally towards L3, with the second last lumbar level being smaller but the last lumbar level being of absolute highest value. Edmondston et al.

(1994) describe for the thoraco-lumbar spine in the elderly, a caudal increase in ventral vertebral body height, except for the mostly constant mid-thoracic region. Ross e/ a/.

(1991) found an increase in anterior vertebral body height, measured in an X-ray study on postmenopausal women, from thoracic levels caudally to L3, with a subsequent slight

decrease for the two last lumbar levels. Jankauskas (1994) in his osteometric study of an

archaeologic sample found a decrease in anterior vertebral body height for level C3 to C6

with an increase in size caudally to L5, mostly similar for females and males. }l4iwte et al.

(1988) list for males a steady increase in anterior vertebral body height caudally, whereas

females reach the highest value in anterior vertebral body height at level L3. In another

radiological study, Hurxthal (1968) found in females an increase in anterior body height

caudally of Th7. Other radiographic studies (Nissan and Gilad, 1984; Gilad and Nissan,

1986) report a decrease followed by an increase for the anterior height of the cervical

vertebrae and similar for the lumbar levels but with an increase from level L4 to L5.

Berry et at. (1987) report a consistent increase caudally in anterior vertebral body height

in the thoracic and lumbar spine. Gallagher et al. (1988) describe an increase in anterior

vertebral body height in a sample of living asymptomatic females from Th3 caudally to

L3 with a slight decrease for L4 and another increase at the last lumbar level. Simiiarly,

Davies et al. Q,989) found in their radiographic study on healthy women that the anterior

vertebral body height increases caudally from Th7 to L4. The anterior vertebral body

height increases caudally in the female cadaver sample, as examined by Aeby (1879),

F. J. Rühli - Osteometric Variation of the Human Spine 35 with the exception of the lower cervical spine and Th6. Cyriax (1920) describes for a

sample of macerated spines of both sexes a consistent increase in anterior vertebral body

height caudally. The thoracic and upper lumbar anterior vertebral body height, according

to the study on cadaver spines by Jacobi (1927), continuously increases caudally.

Tominaga et al. (1995) found mostly an increase in anterior cervical vertebral body

height caudally. In the osteometric study by Marchesi et al. (1988) mainly an increase

caudally in anterior vertebral body height was reported for the mid-thoracic to lumbar

spme.

posterior vertebral body height is strongly correlated with anterior body height

with a correlation coefficient of 0.74 (Ctark et al., 1985). The posterior vertebral body

height shows a steady caudal increase for the whole thoracic spine, with a maximum at

al', Ll and a decrease in size for the rest of the lumbar region (Lanier, 1939; Hermanrt et

1993; Edmondston et al., lgg4). Anderson (1883) describes mostly an increase of

posterior vertebral body height caudally for the thoracic spine and a further increase for

the upper lumbar spine, but a decrease in size for the last three lumbar levels. According

to Minne et at. (I9g8) the posterior vertebral body height increases, as measured in their

study caudally from Th4, in the thoracic spine and reaches in both sexes its highest value

atL3. Hurxthal (1968) found in females an increase in posterior vertebral body height

from Th7 to L3 with a slight decrease for the last two lumbar levels. Putz (1981)

posterior describes a decrease from C3 to C7 with a continuous increase in size for the

the vertebral body height caudally, with a maximum on Ll and a caudally decline within

(1991) lumbar spine. In an X-ray based study on healthy elderly women' Ross e/ al.

found increasing values in posterior vertebral body height from the thoracic spine down in to L2 with a slight decline more caudally. Also Berry et al. (1987) found an increase

36 F. J. Rühti - Osteomelric Variation of the Human Spine posterior vertebral bocly height from upper thoracic level up to L2 with a decrease caudally. Jankauskas (1994) reports, for his historic samples, a decrease in posterior vertebral body height caudally from C3 tlll C7 , with a subsequent increase in size to L 1.

According to him, the lumbar spine shows caudally a decrease in posterior vertebral body height for males but mixed patterns for females. In a study on asymptomatic females,

Gallagher et al, (1988) found an increase in posterior vertebral body height from Th3 to

L3, with a decrease in size for the last two lumbar levels. The same findings were mentioned by Davies et al. (1989), who showed an increase in posterior vertebral body height for their sample of radiologically assessed measurements of female spines caudally of Th7. In the female sample of Aeby (1879) the posterior vertebral body height increases caudally, with the exception of the lower cervical spine. The posterior vertebral body height, as measured by Jacobi (1927) on thoracic and upper lumbar levels, increases caudally with the exception of the mid-thoracic region. Panjabi et al. (1991a; 1991b;

1992) found a decrease ofthe posterior vertebral body height in the upper cervical spine, with a steady increase from the lower cervical spine caudally to level L2.For the cervical spine, Tominaga et al. (1995) found an increase caudally in posterior vertebral body height. Marchesi et al. (1988) report an increase in posterior vertebral body height caudally from the mid-thoracic spine to L3 with a decrease at L4 and L5. Gilad and

Nissan (1984; 1986) found a caudal decrease followed by an increase in the posterior cervical vertebrae height, but on the lumbar level an opposite trend with the highest value forL2.

The particular anatomy of the posterior vertebral surface has been addressed by

Larsen (1985). The foraminae of the basivertebral veins as well as the concave shape of the dorsal part of the lumbar vertebra are highlighted by him. The maximum medial

F. J. Rühli - Osteometric Variation of the Human Spine 37 concave depth was in his sample approximately 0.5 mm for both levels, Ll and L5, respectively. He found no correlation between the posterior vertical lumbar scalloping and posterior vertebral body height or maximum transverse dimension. The scalloping of the lumbar vertebrae is biomechanically explained by Larsen (1985) by various factors, such as axial loading and pressure originating from the cerebrospinal fluid. He explains the development of the lumbar posterior vertebral surface to be a result of its surrounding structure, namely the spinal canal and its contents such as the pressure in the epidural space. The fact that the epidural space changes in its size from cranial to become larger more caudally is another aspect. Furthermore, Larsen (1985) mentions that in cases of narrowing of the spinal cord space the epidural space is altered in form of decreased

content of epidural fat, which may result in an decreased buffer action, which then will

interfere with the posterior surface of the vertebra. The influence of the dural sac and its

contents, according to Larsen (1985), may be more important at the foetal stage, since at

this time the direct physical contact of these two anatomical structures is more intense

than later in life.

The sagittal diameters of the vertebral body, as measured by Lanier (1939) on the

superior and inferior surface level of each vertebra, increase constantly caudally with the

single exception of C7 and L5. Others (Nissan and Gilad, 1984; Gilad and Nissan, 1986)

report for the cervical spine mostly a caudal increase in size and for the lumbar spine a

similar mostly caudal increase for the lower sagittal surface diameter, whereas the upper

sagittal diameter increases only through L3 and decreases further caudal. Katz et al.

(1975) found in an X-ray based study of the cervical spine trends similar as to those

described by Lanier (1939), with C5 having the smallest absolute height, whereas C3

showed the minimal sagittal diameter. For all of the cervical vertebrae, Katz et al. (1975)

F. J. Rühli - Osteometric Variation of the Human Spine 38 found higher values for the sagittal diameter than for the average vertebral body height.

Larsen and Smith (1980b) report in a myelographic study of the lumbar spine an increase of the sagittal vertebral body diameter from Ll to L3 with identical values for the more caudal levels. The results were similar for both sexes (Larsen and Smith, 1980b). Berry et at. (1987) report, with the exception of the mid-thoracic level, a mild increase in the main vertebral body diameters from upper thoracic caudally. Anderson (1883) in an osteometric study found mostly an increase of the sagittal diameter of the vertebral body

caudally. Scoles et al. (1988) report in their study on macerated spines for both sexes in the thoraco-lumbar region a continuous increase in sagittal vertebral body dimension

caudally. Postacchini et al. (1983) found mostly an increase in sagittal and transverse

vertebral body dimension from Ll caudally. Piontek (1973) mentions an increase in

vertebral body dimensions at all levels caudally. This increase was in the cervical spine,

according to him, more prominent in females. For both females and males the sagittal

vertebral body diameter seems to increase caudally, with single exception on a few

selected vertebral levels (Aeby, 1879). Surprisingly, sagittal and transverse cord

diameters, according to Elliot (1945), correlate only vaguely with each other. He also

describes the cervical enlargement of the spinal cord, at level C5 I C6, to be flatter in

sagittal direction, the thoracic to be minimal at level Th6 I Th7 and the lumbar

enlargement at level L5 / S1 to be of small and round shape'

The vertebral body surface area shows in the lower thoracic spine an increase with

a maximum at the second last lumbar level (Shapiro, 1993). This surface area is, again

with the exception of the last lumbar level, well correlated with the body weight, but the

human data in the study by Shapiro (1993) were merged in a sample with great ape. Davis

(1961) examined the relationship between vertebral body area, pedicle dimension and

F. J. Rühti - Osteometric Variation of the Human Spine 39 on transverse processes size in the lumbar spine. He concludes that L4 is larger than L5 average, but it is the other way round for the pedicles. He found no significant correlation between the vertebral body area changes and transverse process size. He explains the caudal transition of the trunk and upper limb weight in the lower lumbar area to be done joints. by lumbo-s acral zygoapophyseal joints but also substantially by ilio-iumbar

The transverse vertebral body diameter, as measured by Lanier (1939) on the inferior surface of the vertebral body, continuously increases caudally, with exceptions of

Th3 to Th6 and at L5. Larsen and Smith (1980b) report a steady increase for the lumbar

the transverse vertebral body dimension caudally. Jankauskas (1994) found for most of cervical levels in males an increase of the transverse vertebral body diameter caudally, a

all of the decrease in the upper thoracic levels and a subsequent increase in size in almost

(1988) more caudal levels. According to him, females show a similar pattem. Scoles et al'

describe for both sexes a steady increase caudally, on the thoraco-lumbar level. Aeby

(1S79) found in his cadaveric sample for both males and females similar trends in

in transverse vertebral body diameters. The transverse diameter decreases caudally both

the upper cervical and thoracic spine, shows an increase in size in the lower cervical and

thoracic as well as the whole lumbar spine (Aeby, 1879). Anderson (1883) describes for

the transverse width of the vertebral body, which was in his osteometric study measured

as the maximal width varying in relative position on each vertebral level, mostly an

increase in size caudally. The only exception in his study was the upper thoracic spine,

which showed a decrease of this measurement caudally from level Th2 to Th5' Cyriax

(1920) found for the transverse vertebral body diameter of a sex pooled sample an

increase in size in the cervical spine, with a slight decrease in the upper thoracic spine and

another increase caudallY.

40 F. J. Rühli - Osteometric Variation of the Human Spine The size of the intervertebral discs, according to radiological studies (Nissan and

Gilad, 1984; Gilad and Nissan, 1986), mostly shows an anterior and posterior increase increases in height in the lumbar segment, whereas no such clear trend is visible for the cervical intervertebral disc anterior and posterior height (Kandziora et a1.,2001). The cervical intervertebral discs are relatively larger than the lumbar ones (Brain, 1948).

Furthermore, Aeby (1879) found that the increase in intervertebral disc size is mainly in the lumbar spine. He provides data on intervertebral disc height for all vertebral levels and both sexes, whereas Tribus and Belanger (2001) only do for the last lumbar one and

Kandziora et at. (2001) for the cervical spine. Jacobi (1927) presents similar results in his sample of cadaver spines from Thl to L3, with the major increase of the intervertebral disc to be found in the upper lumbar spine. Also Piontek and Zaborowski (1973) list normative data on the intervertebral disc height, with mostly an increase caudally for the

cervical spine. Hurxthal (1963) provides in a radiological study on women data for the

lower thoracic and lumbar intervertebral disc heights in normal as well as osteoporotic

individuals. Hurxthal (1968) found an increase caudally. Hasegawa et al. (1995) describe

in their cadaver study of the lumbar spine no increase in intervertebral disc height

caudally.

The maximum spread of the transverse processes increases in the cervical spine

caudally, decreases through the thoracic spine and reaches its smallest size at Thl2.

Finally, it increases again through the lumbar spine and shows its overall highest value at

L3 and L5, respectively (Lanier, i939). Francis (1955) found a decrease of the transverse

process size from Cl to C3 with an increase for the caudal half of the cervical spine'

the Cyriax ( 1920) reports an increase caudally of the total transverse process width within

cervical spine, with a stabilization or clear size decrease within the thoracic spine, and

41 F. J. Rùhti - Osteometric Variation of the Human Spine another strong increase for the lumbar vertebral levels. Panjabi et al. (1991a; 1991b;

1992) report a decrease of the transverse process width for the upper cervical spine with an increase caudally. From Thl caudally, the transverse process width decreases again through Th4, shows caudally a slight increase with a further drop in size at the lowest thoracic levels (Panjabi et al., 1991b). With the exception of L4 this vertebral dimension show an increase in the lumbar spine caudally (Panjabi et a|.,1992).

The spinous process, as measured including the sagittal diameter of the spinal canal in an X-ray study (Nissan and Gilad, 1984), decreases from C3 caudally and increases in size at the caudal half of the cervical spine. According to this particular study, this is not the case for the lumbar level, where there is an opposite trend visible with an initial increase caudally and later decrease in size for the lower lumbar spine.

The normal osseous spinal canal diameters follow mostly a different pattern.

According to Larsen and Smith (1980b) there is no correlation between the main vertebral body dimensions and the spinal canal outline, but there is a correlation between the bony spinal canal and the dural sac size (Larsen and Smith, 1980a).

The sagittal spinal canal dimension shows generally an increase in the cervical segment with its highest dimension on C6, except for C2 (Panjabi et ø1., I99la).In the thoracic spine it shows caudally of Th2 an increase to reach a maximum at Th6, with a subsequent fuither decrease in the lower thoracic spine (Panjabi et al., l99Ib). The two lowest thoracic levels show another increase in size, which continues to Ll (Panjabi et al.,l99lb; Panjabi et al.,1992). From L1 caudally the sagittal dimension decreases to L3 and shows another increase in the last two lumber levels (Panjabi et al., 1992). At L5, with the exception of C2, the overall biggest sagittal spinal canal diameter can be found

(Panjabi et al., l99la; Panjabi et al., I99lb; Panjabi et al.,1992).

F. J. Rühli - Osteometric Variation of the Human Spine 42 increase caudally with relatively The sagittal spinal canal diameter shows a mild - andThll -L2 regions (Lanier' 1939)' and absolutelyhighermeasures forthe c3 c5

a decrease in sagittal spinal canal wolf et at. (1956), based on an X-ray study, describe for the lower cervical levels' Burrows diameters from Cl to C4, *ìrn ,r*ttur values

slight decrease in cervical sagittal spinal (1963) in a similar study, found caudal of C2 a

spinal canal to be shaped like a "triangular canal dimensions, with the osseous cervical much bigger than the sagittal dimension' tube,,, with the interpedicular width being the spinal cord seems to have more than Furthermore, Burrows (1963) mentions that dimension' Dommisse (1974 1975) sufficient space, especially in the transverse the narrowest part in the mid- highlights the fact that the human spinal canal shows involving Th6' Dommisse (1974; 1975) thoracic region, with in most cases particularly

osseous diameters of the spinal canal from describes a decrease in sagittal and transverse

caudally' This most constricted spinal uppef thoracic to the narow zone, with an increase

supply of the spinal cord is also to be least canal region is the region, where the vascular in paraplegia (Dommisse' rich (Dommisse, 1974). This could result in certain instances spinal canal shows an increase in Igl4).Whereas the transverse diameter of the cervical

decrease from c3 to c4 with a mostly stable size caudally, the sagittal dimension shows a

sample (Tominaga et al'' 1995)' In an size caudally, as shown in a mixed-sex cadaveric

(1955), the main diameters of the spinal canal osteometric study conducted by Francis on the sex and populational show an inconsistent size pattern caudally, depending (1955) lists for the sagittal diameter in background of the sample. In general, Francis

upper cervical part with the lower caudal males a decrease in size caudally only for the

values; whereas in females the sagittal half of the cervical spine having roughly similar cervical spine' For the transverse diameter decreases caudally throughout the whole

43 Spine F. J. Rühli - Osteometric Varíation of the Human dimension, Francis (1955) found generally for both sexes a decrease in size in the upper half of the cervical spine and an increase in size for the caudal cervical spine. The minimal sagittal diameter of osseous spinal canal shows, in an osteometric study by

Marchesi et al. (1988), in the mid- / low thoracic and lumbar spine two major values at

Ll and L5, respectively. Aeby (1879) found for both sexes a puzzling pattem of size alterations in sagittal spinal canal size caudally. In general, the canal size decreased caudally in the cervical spine, whereas mostly in the upper thoracic spine this dimension increased and was mostly smaller but stable in size caudally in the lower thoracic spine

(Aeby, 1879). The lumbar spine showed for both sexes caudally an increase followed by a decrease (Aeby, 1879), with males having smaller absolute sagittal lumbar spinal canal diameters than females. Stockdale and Finlay (1980) found in their ultrasound based study a decrease in oblique sagittal lumbar spinal dimension from Ll to L3 with an increase caudally.

The transverse diameter of the spinal canal usually demonstrates two peaks in size, one for the cervical spine and another one for the lumbar region. Obviously, these mark the cervical and lumbar enlargements of the spinal cord, which reflect the increased neural tissue demand for the upper and lower limbs. The cervical enlargement is usually broader than the lumbar one (Elsberg and Dyke, 1934; Elliott, 1945; Maclarnon, 1995).

Magnuson (1944) gives an average size of the osseous transverse spinal canal diameter of

19 mm atL4 and 12 mm atL5. Berry et al. (1987) report for the transverse spinal canal diameter a mild increase caudally, whereas the sagittal diameter did not change from the upper thoracic down to the lower lumbar levels. Aeby (1879) found for both sexes similar trends in transverse spinal canal diameters. He reports a sharp decrease in the most upper cervical spine then mostly a slight increase caudally, with the single exception of the

F. J. Rühli - Osteometric Variation of the Human Spine 4 upper third of the thoracic spine, which shows caudally a decrease in transverse spinal canal size. The transverse diameter of the osseous spinal canal reveals mostly an increase in the cervical spine caudally, with a decrease in size in the upper thoracic spine and generally steady dimensions in the mid-thoracic segment, and a further increase caudally

(Panjabi et al., l99la; Panjabi et a1.,1991b; Panjabi et al., 1992). For the low thoracic and lumbar levels the minimal transverse spinal canal shows, according to a study by

Marchesi et al. (1988), mostly an increase caudally. In their study on recent macerated spines, Postacchini et at. (1983) describe a decrease in mid-sagittal neural canal size from

L1 to L4 with a slight increase for the last lumbar level. The interpedicular distance shows in general the opposite trend. The value of the above mentioned normal limits for the neural canal size in individuals were doubted by Postacchini et al. (1983).

Furthermore, two similar ultrasound based studies on the spinal canal dimensions showed signifrcantly different results (Porter et al.,I978a; Legg and Gibbs, 1984). Nevertheless,

Hinck et at. (1966), based on an X-ray study, provide also normal range values for the interpedicular distance in adults. This particular landmark demonstrates an increase in the middle cervical spine caudally and decreases towards mid-thoracic spine, with a final continuous increase from mid-thoracic to low lumbar levels. Already Elsberg and Dyke

(1934) defined and reported normal values for the interpedicular distance of all vertebral

levels, as measured on conventional X-ray films. They describe a similar pattern of

normal interpedicular morphology, as did Hinck et al. (1966). Furthermore, Gepstein e/

al. (1991) found an increase in lumbar interpedicular distance caudally and also for the

mid-cervical spine. According to Eisenstein (1,977), the normal interpedicular diameter

measures 23 mm in the lumbar spine and shows no noteworthy variation within the

lumbar levels.

F. J. Rühti - Osteometric Variation of the Human Spine 45 A decrease in spinal canal dimension from Ll caudally was reported in another ultrasound study by Macdonald et al. (1984), with a slight increase for the last lumbar level. The values were slightly different in symptomatic individuals with another decrease in size for the last lumbar level (Stockdale and Finlay, 1980). In a similar study, Legg and

Gibbs (1984) found mostly a decrease for the lumbar spinal canal dimension caudally.

Williams (1975) explored the pathologic narrow lumbar spinal canal relative to the vertebral body size at the same level, with a ratio of 116 or 1/6.5 to be defined as being pathologic. The major spinal canal dimensions were investigated by Scoles et al. (1988) for both sexes on selected thoraco-lumbar levels in a sample of macerated spines. They describe an increase in sagittal diameter caudally in the thoracic region, followed by a decrease in the upper lumbar and another increase in the lower lumbar spinal levels. The transverse diameter, as described in the study by Scoles et al. (Scoles et aL.,1988) shows also similar for both sexes a decrease in size in the upper thoracic spine with an increase caudally in the lower thoracic and lumbar spine. For the lumbar spinal canal, Huizinga et al. (1952) found an increase in interpedicular width only for the last lumbar level, whereas the antero-posterior spinal canal dimension shows a decrease from Ll to L3 with a slight increase further caudal. A significant relation between these two vertebral canal dimensions are found only for L3 and L4 (Huizinga et al., 1952). Kikuchi et al. (1977) describe for the sagittal diameter of the lumbar spinal canal a decrease caudally of Ll with a subsequent increase for the last two lumbar levels, whereas the interpedicular canal shows a steady increase caudally. Larsen and Smith (1980a) found a decrease in size for the mean sagittal diameter from Ll to L4 with an increase for the last lumbar level, whereas the transverse diameter showed a steady decrease in size in the lumbar spine

caudally. The lumbar subarachnoid space, according to them, was the smallest in sagittal

F. J. Rühli - Osteometríc Variation of the Human Spine 46 direction at L4. As Larsen and Smith (1980a) pointed out, the subarachnoid space consists mostly of blood vessels and loose tissue and measures I to 3 mm. Clark et al.

(1985) mention a significant correlation of the two main spinal canal diameters at thoracic and lumbar levels in two historic samples.

Critical values of spinal canal dimensions are reported in various studies. Epstein et al. (1964) mention a sagittal diameter of less than 13 mm to be of pathologic value. If so, the condition was often accompanied by short and bulky pedicles and massive neural arches (Epstein et al., 1964). Similar critical values for X-ray measurements are mentioned by Wolf et al. (1956). In a clinical study, Porter et al. (1987) group patients in two samples, based on their 15" oblique sagiffal spinal canal diameter on Ll, with the very narrow ones being below 14.1 mm and the very wide ones being above 15.8 mm.

The averages for these two particular groups were 13.8 mm and 16.4 mm, respectively. In other ultrasound based studies, Porter et al. (1978b; 1980) report data of the normal sagittal lumbar spinal canal diameter. They found a decrease in 15o oblique size from Ll to L4 with another increase atL5, consistent for both sexes, but in general slightly bigger for females. They mention the cut-off point in oblique sagittal neural canal diameter for people at clinical risk as being 14 mm. Eisenstein (1977) declares as lowerlimits of the interpedicular distance a width of 18 mm, whereas the normal width, according to him, seems to be 23 mm. Eisenstein (1980) further reports in another study that the trefoil shape of the lumbar spinal canal is mainly caused not by osteophytic overgrowth but rather by a local thickening of the laminae. \ù/olf e/ al. (1956) mention a minimal sagittal diameter of the cervical spinal canal of 10 mm, based on X-ray assessments, to avoid clinical symptoms in form of spinal cord compression. Eisenstein (1977) states for the sagittal lumbar vertebral foramen diameter values of 13 mm and 16 mm, respectively.

F. J. Rühli - Osteometric Varialion of the Human Spine 47 Larmon (1944) mentions as average size for the intervertebral foramen width in cadavers of 7 mm, similar to the mid-sagittal diameter of the vertebral canal, whereas the transverse diameter of the vertebral canal was in his sample on average 12 mm. Kirkaldy-

Willis et al. (1982) list 4 mm as a borderline for the lumbar intervertebral foramen width as measured on CT scans. Hasegawa et al. (1995) declare a posterior disc height of 4 mm and a foraminal height of 15 mm as crucial minimal limits.

The cross-sectional area of the lumbar spine was calculated in a cadaver study by

Hasue et al. (1983). For the osseous and non-osseous dimensions of the spinal canal in males, the largest value was found atL5, with the smallest being at mid-lumbar, whereas no such trend was found in females. The mean areas of the neural tissue become smaller in caudal direction for both sexes, with the single exception of an increase at L5 in females (Hasue et a1.,1983; Kikuchi et a1.,1984). According to Hasue et al. (1983) and

Kikuchi et al. (1984), males have larger osseous and non-osseous dimensions, except at

L5, but have smaller neural tissue sizes than females. Similar trends can be found for the relation between spinal nerve and the osseous and non-osseous intervertebral foramen size (Hasue et al., 1983; Kikuchi et al., 1984). The lumbar spinal canal, at least as reported for symptomatic subjects (Porter et a1.,1978b), shows side differences in its 15o oblique diameter, varying between 0.4 mm at L1 and 0.7 mm atL4.

The pedicle size increases in humans in the lower thoracic spine caudally, decreases slightly in the upper lumbar part and shows another but even stronger increase towards the caudal end of the lumbar spine (Shapiro, 1993). The most striking enlargement of pedicle sizes occurs between the second last and the final lumbar vertebra with an average increase of 73o/o (Shapiro, 1993). Human pedicle size, as pointed out by

Shapiro (1993), is correlated with body size for most lower back levels; unlike pedicle

F. J. Rühli - Osteometric Variation of the Human Spine 48 shape, which is defined as the ratio of pedicle width to pedicle length. The pedicle height, as described for a sex pooled cadaveric sample by Tominaga et al. (1995), shows an increase from C3 to C4, with a decrease caudally and another increase in the last cervical level. Zindrick et al. (1987) found, in their radiological cadaver study, for the pedicle length an increase in the thoracic spine to Thll and a decrease caudally. Krag et al.

(1988) in a similar study, report that the pedicle length decreases from Th9 caudally.

Misenhimer et al. (1989) describe a decrease in pedicle height caudally for the upper thoracic levels with an increase through Thl2. The lumbar levels show a similar pattern, as do the thoracic, with a decrease in size for the upper part and an increase for the lower levels (Misenhimer et al., 1989). Furthermore, Misenhimer et al. (1989) state that the thoraco-lumbar pedicles have a teardrop-shape with the widest part in the inferior half.

Saillant (1916) describes an increase in pedicle height from C7 caudally, most prominent for the upper thoracic and the most lower thoracic spine, with a caudal decrease in size for the majority of the lumbar levels. Ebraheim et al. (1997) found predominantly a slight decrease caudally in pedicle height for the mid-cervical spine in both sexes. For the cervical pedicle width Karaikovic et al. (1997) found generally an increase in size caudally. In a study on the pedicle dimensions in an Indian population, Mitra et al. (2002) report a decrease in pedicle length from Ll to L4 with a size increase at L5, with females having non-significantly larger values in general. Olsewski et al. (1990) found for males a decrease in size of the pedicle height from Ll to L2 with an increase caudally. Females showed a decrease caudally of Ll with only an increase in size for the last lumbar level

(Olsewski et al., 1990). The pedicle angle, as measured in relation to the sagittal axis, increases especially in the lower lumbar spine (Krag et al., 1988). Scoles et al. (1988) report for both sexes an increase in maximum pedicle diameter caudally for most of the

F. J. Rühli - Osteometric Variation of the Human Spine 49 thoracic levels, with a decrease in size in the upper lumbar region. They found the overall largest values for the maximum pedicle diameter on the lowest lumbar level. Vaccaro et al. (1995) describe mostly a slight increase in mid-lower thoracic pedicle height caudally, similar to the frndings by Hou et al. (1993). Vaccaro et al. (1995), furthermore, describe a decrease caudally in the lumbar spine, with a sharp increase in pedicle height for the last lumbar level. The pedicle height, as measured on all levels and bilaterally by Panjabi et al. (l99la; 1991b; 1992), shows in general a decrease in size caudally in the upper cervical, mid-thoracic and upper- / mid-lumbar spine, whereas an increase caudally can be found in the other spinal regions. The highest values are reported for level L5 (Panjabi et al.,1992).

The particular anatomy of the dural sac, the shape of the dural sheath at lumbar levels and the lumbo-sacral nerve roots, have been widely addressed by Salamon et al.

(1966). The size of the subarachnoid space in the lumbar spine of symptomatic and asymptomatic individuals has also already been addressed (Larsen and Smith, 1980a).

The lordosis of the cervical spine is, according to Jankauskas (1994), only caused by the intervertebral discs, whereas the lumbar one is formed by the discs as well as the vertebral bodies' shape. The lumbar lordosis was to be found more prominent in females and in individuals of greater body weight (Murrie et a1.,2003).

The size of lumbar intervertebral foramen is usually the biggest for L5 / S1, whereas Ll I L2 have the smallest area (Stephens e/ al., l99l).Putti (1927) describes the opposite, with the L5lS I intervertebral foramen being the smallest and lists as a rule that the more cranially located, the bigger the lumbar foramen is supposed to be. On the other hand, Putti (1927) mentions the contrary for the nerve root size, with L5 being the largest, and the more cranial one being smaller in size. Kirkaldy-Willis e/ al. (1982) list,

F. J. Rühli - Osteometric Yaríation of the Human Spine 50 independent of the vertebral level, an average of 4 mm as a borderline for the intervertebral foramen width, as measured by CT. Ebraheim et al. (1996) reported the dimensions of the cervical intervertebral foramen in cadaver and macerated specimens.

They divided the cervical intervertebral foramen in three parts, a medial pedicle section, the middle section next to the foramen transversarium, and finally, a most lateral part.

Except for the first level at C2 I C3, all other cervical levels showed, according to

Ebraheim et al. (1996) an increase in size caudally. The minimum intervertebral foramen width was 1-2 mm at all levels.

The weight of the dry human spine differs by region and sex. According to Trotter and Hixon (1974), the cervical, thoracic and lumbar vertebrae of White adults weigh approximately 53 g, 131 g, II2 g for males and for females 39 g, 98 g, and 8l g, respectively. The relative weight of the axial postcranial skeleton, consisting for this analysis of the vertebral column, ribs and sternum, decreases mostly in adulthood in both sexes, and is on average 18% of the total adult skeleton weight (Trotter and Hixon, 1974).

Additionally, Trotter and Hixon (1974) found that males show higher mean bone densities in all spinal parts.

Impact of osteometric research of the human spine

Spinal osteometric data can be applied for various purposes. They help e.g., to

estimate stature, since the size, weight and volume of the spine are usually correlated with individual height in humans (Hasebe, 1913; Martin, 1928; Latimer, 1950; Martin and

Saller, 1957; Fully and Pineau, 1960; Tibbetts, 1981; Jason and Taylor, 1995).

Nevertheless, in the osteometric study by Berry et al. (1987) the size of the combined vertebral body heights did not correlate with the individual body height at autopsy.

F. J. Rühli - Osteometric Variation of the Human Spine 5L Furthermore, morphometric studies of the osseous vertebral column help to define gold-standards for subsequent clinical applications (Saillant, 1976; Kikuchi et al., 1977; postacchini et a1.,1983; Nissan and Gilad, 1984; Gilad and Nissan, 1986; Berry et al',

1988; 1987; Ziñ¡1ck et al., 1987;Y:rag et al., 1988; Marchesi et a1.,1988; Scoles et al',

Misenhimer et a1.,1989; Olsewski et a\.,1990; Black et a\.,1991; Panjabi et al',l99ta;

1993; Panlabi et al., 1991b; Panjabi et al., 1992; ]ËIletmafIrl et al., 1993; IJou et al., vaccaro et al., 1995; Xu et al., 1995; Kothe et al., 1996; Ebraheim et al', 1997;

Karaikovic et al., i997; Mitra et a1.,2002) or they can proof the suitability of animal

et al', models in relation to the human spinal dimensions (Cotterill et al.,1986;Tomitaga

vertebral 1995). panjabi et ql. (1992) declare their study on the three-dimensional

or in morphometry to be useful as a "blueprint", which can be implied in clinical issues

(1988) the fact mathematical analysis of the spine. Furthermore, Scoles et al' emphasize

orthopaedic that the knowledge of spinal morphometry is still limited, despite its need for by implant assessments. For example, Scoles et al. (1988) and a similar study undertaken

Berry et al. (1987) disagree on the minimum pedicle dimensions, which would have a

crucial impact on the use of transpedicular f,rxation screws. In their study on Indian published populations, Mitra et al. (2002) also found pedicle values different from earlier be used in ones, which led them e.g., to recommend specific screw dimensions to

chirurgical approaches. Similar observations are reported for the non-'White sample

pedicle dimensions than examined by Hou et at. (1993), which showed in general smaller

earlier reported standards.

A list of performed earlier major osteometric studies could be found in Table 1,

are listed in whereas morphological studies on cadaveric samples and living individuals

Table2.

52 F. J. Rühti - Osteometric Variation of the Human Spine Table I Osteometric studies of recent and historic spine samples

.Reference Time Period N Method Spinal region

Aeby (1879) Late 19ù century 13 Scale All

Amonoo-Kuofi (1985) Recent 92 Caliper Lumbar

Anderson (1883) I-ate 196 cenh¡ry 53 Ruler At1

Berry et al. (1987) Late 19û / Early 20' 1 Caliper? All cetrtury

Boszczyk et al. (2001) Recent 106 Anthropometer All

Clark et al. (1985) 10*-13" century ?t9s Caliper Al1

Cotterill et al. (1986) Recent l0 Caliper Th6, ThI2 and L3

Cwirko-Godycki and Swedborg l3* century 48 Caliper CIIC2 (re7'7)

Cyriax (1920) Early 20* century Ca'10 ? All

Davis (1961) Mid 20ù century 201 Caliper A1l

Dommisse Q97a;1975) Recent 6 /25 Caliper Thoracic and lumba¡

Ebraheimetc/. (1996) Recent 443 Caliper Lumbar

Ebraheimelai. (1997) Recent 40 Caliper Cewical

Eisenstein (1977) Late 19* / Early 20" 338 Caliper? L3/LA cenhrry

Ericksen (1976) L^te 19ú I Early 20* 34 Caliper / clay casts Lumbar century

Francis (1955) Mid 20" century? 284 Caliper Cervical

Frey (1929) Early 20* cenhrry 150 Measurement tape All

,| Fully and Pineau (1960) Mid 20^ centruy r64 All

Gepstein et al. (1991) Recent 54 Caliper Cewical and lumba¡

Hasebe (1913) Early 20" century 30 Measuremeot tape All

Hou et al. (1993) Recent 40 Caliper Th9-L5

Huizinga et al. (1952) 19* century 5l Caliper Lumbar

Iacobi (1927) Early 20* century t02 Ruler Thl-L3

Jankauskas (1994) l'/ 2"oMillenniumA.D. 539 Caliper All

Kaliszewska (1966) 12* century I Caliper AI

F. J. Rühli - Osteometric Variation of the Human Spine 53 Kanziora et al. (2001) Recent 20 Digital ruler Cervical

Karaikovic et al. (1997) Recent 53 Caliper / CT Cervcical

Kikuchi et al. (1977) Recent 80 Caliper Lumba¡ lanier (1939) Recent 30 Caliper T\2,T\7, Thl2 and lumbar

I-ee et al. (1995) Recent 90 Caliper Lumba¡

Marchesi et al. (1988) Recent 33 Caliper? / X-ray Th6 - Ls

Piontek (1973); Piontek and 12-14ú century / l4-186 4rl 50 Caliper Cervical Budzynska (1972); Piontek and cenh¡ry Zaborowski (1973)

Porter and Pavitt (1987) Anglo-Saxon and Roman- Photographic Lumbar British

Postacchini ¿f ø/. (1983) Recent? 12l Caliper Lumbar

Present study Since Late UPPer 348 Caliper c3, c7, Thl, Th6, Thl0, Paleolithic to mid 20'o Ll, L5 centurY

Putz (1981) Recent 66? Scales / Goniometer Alt ')) Ravenel (1877) l,ate 19ú century Scale Alt

Rosenberg (1899) Late 19* century 5 Compass I-ow thoracic and lumbar

thoracic and Scoles e/ a/. (1988) Late 196 / Early 20^ 50 Caliper Selected century lumba¡ levels

and lumbar Shapiro (1993) Recent 42 Low ttroracic

,) Stefl

Swedborg (1974) 10*-12"century 9l? Caliper All

and lumbar Tatarek (200I) Prehistoric / ¡ecent 90 Caliper Thoracic

Thomson (1913) Early 20ù century 6 Caliper? All

Lumbar Todd and Pyle (1928b) Early 20' 59 Surface drawing century

Tomin¿g¿ et al. (1995) Recent 6 Caliper Cervical

Wetzel (19I0) Late I 9" / Early 20' 16 ? All century?

,| Wood-Jones (1938) Early 206century? ? ClIC2

Xn et al. (1995) Recent 56 Caliper C7

F, J. Rühlí - Osteometríc Variation of the Human Spine 54 TabIe 2: Morphological studies of spines in living people and cadavers

Reference Material N Method Type Spinal Region

Adams af a/. (1994) Cadavers l9 Biomschanical Dynamic Lumba¡

Banta e/ a/. (1989) Cadaven l6 Caliper / X-ray Static Th6 - L5

Daviesefai. (1989) Living, l9l X-ray Static ^t"h't -IA asymptomatic lryOmen

Diacinti et al. (1995) Living, 126 X-ray Static Thoracic and lumbar asymptomatic v/omen

Dommisse (1974; 19'75) Living, 50 X-ray St¿tic Tho¡acic and lumbar asymptomatic

Drinkall et al. (1984) Asymptomatic and 386 Ultrasound St¿tic Lumbar symptomatic

Ebr¿heim et al. (1996) Cadaven 14 Caliper Static Cervical

Edmonston e, al. (1994) Cadavers of elderþ l8 CT Static Thoracic and lumba¡ people

Elsberg and Dyke (1934) Asymptomatic / 100 / X-ray Static All symfrtomatic 86

Fujiwara et al. Q00I) Cadavers 39 CT lbiomechanical Dynamic Lumbar

Gallagher and Hedlund Living, 150 X-ray Static Th3 - L5 (1e88) asymptomatic v/omgn

Gozdziewski et al. (1976) Living, 77 6 Antlropometric Static Thoracolumba¡ asymptomatic

Harrington et al. (200I) Living,symptomatic'72 CT St¿tic L4 / L5 only and asymptomatic

Hasegawa et al. (1995) Cadaven l8 Photographic Static Lumba¡ measuremeûts

Hedlund and Gallagher Living, symptomatic 153 X-raY Static Thoracic and lumba¡ (1e88) women

Herma¡n et al. (1993) Living, I 13 X-ray Static Mid-thoracic and lumba¡ asymptomatic

Hinck er al. (1966) Living, no obvious l2l X-raY Static All pathology

Homer (1854) Cadaver / living, 4?l Ruler? Static / AI asymptomatic t? Dynamic

Humphreys et al. (1998) Living, 43 MRI Static Cervical asymptomatic and symptomatic F. J. Rühli - Osteometric Variation of the Human Spine 55 Hunthal (1968) Living, 20 X-ray Static Th7 - L5 asymptomatic women lnutusa et a/. (1996) Cadavers t7 CT Static and Lumbar dynamic

Jason and Taylor (1995) Cadaven r67 Flexible ruler St¿tic All

Katz et al. (1975) Living, 6l X-ray Static Cervical asymptomatic

lovels Kothe et al. (1996) Cadaven l4 X.ray Static Selected thoracic

IGag et a/. (1988) Living, symptomatic 41 CT Static T9.L5

Larsen and Smith (1980a; Symptomatic and 83 X-ny / Static Lumbar 1980b) asymptomatic Myelography

Legg and Gibbs (1984) Living, 50 Ultrasound Static Lumba¡ asymptomatic males

L,t el al, (2000) Cadavers l6 Computer-assisted Dynamic Cervical photogra.Phic simulation

Macdonald e¡ al. (1984) Living,symptomatic 204 lJltrasound Static Lumbar and asymptomatic

Mapuson (1944) Cadaven 10 Caliper? Static Lumbar

Lumbar Mayoux-Benhamou e/ a/. Cadavers 7 Caliper / Cast Dynamic (1e8e)

low-thoracic and Mime e/ c/. (1988) Living, il0 X-ray Static Mid- and asymptomatic lumbar

lumbar Misenhimer e/ a/. (1989) Cadavers 6 Caliper / CT Static Thoracic and

Mífaet al. (2002) Cadavers 20 Caliper, X-ray and Static Lumbar CT

Nissan and Gilad (1984; Living, t57 X-ray Static Cervical and lumba¡ 1986) asymptomatic

Nowicki et al. (1996) Cadavers 3l CTlMRI Dynamic Lumbar

Olsewski et al. (1990) Cadavers, living 100 Czliper lX-ray, CT Static Lumba¡ symptomatic

Lumbar Panjabi et a/. (1983) Cadaven l2? Biomechanical Dynamic

Cervical Parjabi et al. (l99|a) Cadavers t2 Biomechanical Dynamic

Thoracic Panjabi e/ ci. (199lb) Cadavers t2 Biomechanical Dynamic

Lumbar Piera et øi. (1988) Living, symptomatic 2t5 X-ray Static

Piontek and Zabo¡owski Living patients 185 X-ray Static Cewical (1e73)

F. J, Ri)hli - Osteometric Variation of the Human Spine 56 Porter et al. (1978a) Living, 273 Ultrasor¡nd Static Lumbar asymptomatic and symptomatic

Porter et a/. (1980) Living, 550? Ultrasound Static Lumbar asymptomatic and symptomatic

and lumbar Ross et a/. (1991) Living, 1098 X-raY Static Thoracic asymptomatic women

lumbar Saillant (197ó) Cadavers 35 Caliper? Static Thoracic and Lumbar Schmid ef a/. (1999) Living, 12 MRI (open) Dynamic asymptomatic

Lumbar Stephens et al. (1991) Cadavers 20 Molding technique Static and X-ray

Lumbar Stockdale and Finlay Asymptomatic and +/' Ultrasound Static (1e80) symptomatic 100

All Tibbetts (198 1) Cadavers 200 CaliPer Static

Lumba¡ ullrich ef a/. (1980) Living, 60 CT Static asymptomatic

Mid- / Lower thor¿cic Vaccaro et al. (1995) Cadavers, 36 Caliper, CT Static asymFtomatic patients

Van Schaik ¿l a/. (1985) Living,symptomatic 123 CT Static L3.L5

lumbar Weisz and Lee (1983) Living, symptomatic 75 cr Static Low

Lumba¡ \Vildermuth et al. (1998) Living, symptomatic 30 MRI, MYelograPhY Dynamic

Williams (1975) Living, symptomatic 100 MyelograPhY Static L3-L5

Cervical Wolf ¿l ¿i. (1956) Living, 200 X-raY Static asymptomatic

Ztndrìcket al. (1987) Cadavers 522- X-ray and CT Static Thoracic and lumba¡ 628

57 F. J. Rühti - Osteometric Variatíon of the Human Spine The study of the bony outline of the lumbar spinal canal allows drawing limited conclusions about the size of the dural sac in the particular individual, since these two measurements are mostly well correlated (Larsen and Smith, 1980a). The measured osseous spinal canal dimensions may at least partially reflect the outline of its neural content. Spinal canal dimensions correlate well with spinal cord size in primates

(Maclarnon, 1995; Maclarnon,1996a). Maclarnon (1995) found that the white matter size, unlike the grey matter dimensions of the spinal cord, correlates with the osseous spinal canal dimensions. Maclarnon (1995) links partially the found differences of the spinal canals in various primates with their particular fore- and hind-limb innervation.

Therefore, if one hnds an alteration of the osseous spinal outline this may have various functional implications as well. Maclarnon (1995) interprets the larger dimensions in more dominant limbs is more likely due to more or thicker nervous fibres instead of higher numbers of nerve cells. Whatever the underlying factor, such as increased myelination of fibres or more branched nervous fibres, an apparent increase of neural transmission speed of more developed limb innervation is visible in an altered white matter pattern and, following, the osseous dimensions of the vertebral column

(Maclamon, 1995). Maclarnon (1995) also found that any increased neuronal demand in a limb is more reflected in a more prominent development, within the spinal cord white matter, of the dorsal rather than the latero-ventral columns. This finding was interpreted by Maclarnon (1995) as a reflection of a possible higher proprioceptive demand rather than in numbers of motor neurons. Any increased neuronal supply in a limb seems, therefore, to be mainly influencing the sensory and ltbrous part of the spinal cord and, therefore, be also present in an increased number of sensory neurons, to be found in the dorsal root ganglion that plays a crucial part in the etiology of lower back pain not in an

F. J. Rühli - Osteometric Variation of the Human Spine 58 altered motor neuronal supply. Lassek and Rasmussen (1938) interpret the high variability of white spinal cord matter cross-sectional area as a result of inter-individual

differences in f,rbre size and number. Spinal cord weight shows no clear correlation with

locomotion pattern at least in primates (Maclarnon, 1996b). Surprisingly, the white

matter seems to be more sensitive than the grey matter to such locomotive alterations.

Humans have a higher amount of such white matter in their lumbar spinal enlargement

than predicted for their body weight (Maclarnon, 1996b). It is well known that the

number of neurons and the size of central nervous tissue decrease with age (Dunn, 1912)-

Earlier studies also showed a notable decrease of the myelinated spinal nerve root fibres

with age (Dunn, I9l2; Corbin and Gardner, 1937). Furthermore, it is also well known

number that muscle mass, represented by robusticity in skeletal specimens, influences the

and size of nerve fibres (Dunn, lgl2). For example, the size of the spinal cord and brain

parts are positively influenced in growing cats undergoing physical training (Agduhr,

Dunn lglT), at least for the regions that are innervating the trained muscles. Additionally,

(lgl¿) reports for albino rats a correlation of size and number of cervical nerve root fibres

with increased weight, but also with age to a certain point, before the senile hbre size

and size decrease start. FurtheÍnore, he suggests a correlation between nerve root calibre

of tissue innervated. Corbin and Gardner (1937) also found such a clear correlation

between muscle mass and number of ventral root fibres in the spinal column of selected

individuals.

Spinal pathologíes - especially lower backpain

The vulnerability of the human back produces various neurological and

orthopaedic pathologic conditions (Simmonds, 1903; Bailey and Casamajor, 1911;

59 F. J. Rühli - Osteometric Variation of the Human Spine Willis, 1924; Jacobi, 1927; Putti, 1927; Willis, 1929; Blumensaat and Clasing, 1932;

Philipp, 1932; Samuel, 1932; Junghanns, 1933; Larmon, 1944; Magnuson, 1944;Brain,

1948; Pallis et a\.,1954; Verbiest, 1954; Gill and White, 1955; Nathan et aL.,1960; Roaf,

1960; Epsteinet al., 1962; Burrows, 1963; Epsteinet al., 1964; Hurxthal, 1968; Jones

and Thomson, 1968; Dommisse, 1974; Swedborg,1974; Veleanu, 1975;MacGibbon and

Farfan, 1979;Cinc et al., 1980; Park, 1980; Crock, 1981; Kirkaldy-Willis et al., 1982;

Dorwart et a\.,1983; Ogino et a\.,1983; Louis, 1985; Resnick, 1985; Gaskill et al., l99I;

Jankauskas, 1992; An and Glover, 1994). Furthermore, one has to remember that the

occurrence of some spinal pathologies are inter-correlated with each other (Swedborg,

re74).

Low back pain and other severe clinical symptoms, such as radiculopathy, are

extremely common (Brown, 1975; Kelsey and'White, 1980; Macdonald et al., 1984;

Hartvigsen et a1.,2001; Stebler et a1.,2001) and cause enorTnous socio-economic costs in

modern societies (Macdonald et al., 1984; Gaskill et al., 1991; Maniadakis and Gray,

2000). Therefore, approaches to determine their possible etiologies are numerous (Bailey

and Casamajor, 1911; Willis, 1924;Putti, 1927; Willis, 1929;Blumensaat and Clasing,

1932; PhlIipp, 1932; Mixter and Barr, 1934; Lartnon, 1944; Magnuson, 1944; Brain,

L948;Httizinga et al., 1952; Pallis et a1.,1954; Gill and White, 1955; Epstein et al., 1962;

Burrows, 1963; Epstein et a\.,1964; Nachemson, 1966; Salamon et aL.,1966; Jones and

Thomson, 1968; Brown, 1975; Ramani, 1976; Eisenstein, 1977; Kikuchi et al., 1977;

Porter et al., I978a; MacGibbon and Farfan, 1979; Nachemson et al.,1979; Cäc et al.,

1980; Eisenstein, 1980; Larsen and Smith, 1980a; Larsen and Smith, 1980b; Porter et al.,

1980; Crock, 1981; Hasue et a\.,1983; Ogino et a\.,1983; Panjabi et a|.,1983; Weisz and

Lee, 1983; Jungers, 1984; Kikuchi et al., 1984; Macdonald et aL.,1984; Rydevik et al.,

F. J. Rühti - Osteometric Variation of the Human Spine 60 1984; Vanderlinden, 1984; Clark et a|.,1985; Weinstein, 1986; Heliövaara,1987; Porter et al., 1987; Porter and Pavitt, 1987; Rauschning, 1987; Hoyland et al., 1989; Mayoux-

Benhamou et al., 1989; Yoo et al., 1992; Yoshida et al., 1992; Ebraheim et al., 1996;

Nowicki et al., 1996; Leboeuf-Yde et al., 1997; Schmrd et al., 1999; Fujiwara et al.,

200I; Harrington et al., 2001; Hartvigsen et al., 200I; Cinotti et al., 2002; Al Faraj and

Al Mutairi, 2003; Murrie et a|.,2003) and the clinical and diagnostic impact of low back pain has been described for more than one hundred years, as already reviewed earlier

(Dyck, 1984; Rüttimann, 1990; Wiltse,l99l; An and Glover, 1994).

Various radiological techniques, such as conventional X-ray, ultrasound, myelography or CT-scanning, can be used in clinical situations to address the size of the neural pathways and vertebral bodies (Burrows, 1963; Hun

Ramani, 1976;Porter et al., 1978a; Porter et a1.,1978b; Larsen and Smith, 1980a; Park,

1980; Porter et al., 1980; Stockdale and Finlay, 1980; Hibberl et al., 1981a; Kirkaldy-

Willis et al., 1982; Legg, 1982; Weisz and Lee, 1983; Drinkall et al., 1984; Legg and

Gibbs, 1984; Macdonald et al., 1984; Bolender et al., 1985; Gallagher et al., 1988;

Hedlund and Gallagher, 1988; Minne et al., 1988; Davies et al., 1989; Schmid et al.,

1999) or pedicle dimensions (Zindrick et al., 1987; Krag et al., 1988; Marchesi et al.,

1988; Banta et a|.,1989; Misenhimer et al.,1989; Olsewski et al.,1990; Hou et a\.,1993;

Vaccaro et a1.,1995; Kothe et al., 1996; Ebrahetm et al., 1997; Karaikovic et al., 1997;

Mitra et al., 2002). Lumbar spine imaging counts for approximately 4o/o of all X-ray facility workloads (Park, 1980), with a lot of them dealing with lower back pain issues.

Imaging data, gained even with most sophisticated techniques such as advanced

CT-scanning and MRI, differ slightly from data obtained in situ. Black et al. (199I) and

Hermann et al. (1993) remind that morphological measurements obtained from

F. J. Rühli - Osteometric Variation of the Human Spine 6L conventional radiographs may differ depending on the positioning of measuring landmarks. Jones and Thomson (1968) recommend, based on their experience in clinical cases, the use of the vertebral canal to vertebral body ratio in plain X-rays as a supplementary aid. This récommendation was followed in various studies e.g., in a myelographic study on the narow lumbar spinal canal by Williams (1975) or in an osteometric study by Kikuchi et al. (1977). No difference between pedicle measurements obtained by either conventional X-ray or CT scanning was found by Zindrick et al.

(1987), a statement mostly supported by Krag et al. (1988) too. Karaikovic et al. (1997) also mentioned that there is no relevant difference between caliper based measurements and CT data of the same spinal structure. Mitra et al. (2002) found slightly different values of various pedicle dimensions for X-ray and CT-scanning in comparison to direct measurements, similar to Misenhimer et al. (1989); whereas Marchesi et al. (1988) did not found any signihcant difference. Olsewski e/ al. (1990) report mostly significant differences between osteometric and X-ray measurements of various lumbar pedicle dimensions.

Spinal stenosis is a clinical syndrome, which originates from a narrowing of the spinal canal, the lateral recess or the neural foramen as a result of bony and / or soft tissue alterations (Bailey and Casamajor, 1911; Putti, 1927;Larmon, 1944; Magnuson, 1944;

Brain, 1948; Pallis et al., 1954; Verbiest, 1954; Epstein et al., 1962; Burrows, 1963;

Epstein et a\.,1964; AmoIdi et a\.,1976; Kikuchi et a|.,1977; Porter et a|.,1978a; Porter et a\.,1980; Crock, 1981; Kirkaldy-Willis ¿/ al.,1982; Dorwart et a|.,1983; Hasue et al.,

1983; Ogino et al., 1983; Postacchini et al., 1983; Weisz arrd Lee, 1983; Bose and

Balasubramaniam, 1984; Kikuchi et aL.,1984; Rydevik et a|.,1984; Vanderlinden, 1984;

F. J. Rühli - Osteometric Varialion of the Human Spine 62 Bolender et a\.,1985; Rauschning, 1987; Lee et a\.,1988; Hoylarrd et al', 1989; An and

Glover, 1994;Nowicki et al.,1996; Fujiwara et aL.,2001)'

Radiological abnormalities of the cervical spine can be found frequently and in general more coÍrmonly in men (Pallis et al., 1954). Roughly, 75Yo of the individuals aged 50 years and above show a narrowing of the spinal canal due to various underlying

conditions, such as osteophles or vertebral subluxation. Surprisingly, in such a sample of

individuals without neurological symptoms, a similar fraction of adults showed

radiological signs of foraminal narrowing and even more had signs of a narrowed

intervertebral disc space or marginal osteophytes on the anterior vertebral body border

(pallis et al.,lg54).In an unselected sample of individuals who underwent myelographic

imaging, Williams (1975) found a total of 3o/o narïow lumbar spinal canals. In a Danish

longitudinal study assessment, investigated by Hartvigsen et al. (2001), it was found that

heavy work load is important for the occurrence of low back pain and sedentary work acts

protectively. Hartvigsen et al. (2001) discussed this result with regard to the "healtþ

worker effect", which confuses findings of cross-sectional studies on the prevalence of

lower back pain, due to the self-selection process of healthier individuals remaining in

their job; a bias occurring in form of a migration between possible exposure groups.

Heliövaara (1987), as already mentioned above, found a correlation between herniated

lumbar intervertebral disc and body height as well as body mass in males'

Classification of the spinal stenosis etiology usually differentiates between the

congenital-developmental and the acquired forms (Arnoldi et al.,1976)' Most patients are

approximately 35-65 years old, with the majority being over 50 years, and express

various clinical symptoms and signs such as senso-motoric defects, dysfunction of the

bladder, gait instability and radicular pain. A clinical sample of a general practice in rural

63 F. J. Rühti - Osteometric Variation of the Human Spine England showed a slightly higher, but statistically not significant, frequency of low back pain in males with 54% of all 193 cases and a mean age for patients of 45 years (Drinkall et al., 1984). In a sample investigated by Harrington et al. (2001) the average age of symptomatic patients was 43 years for men and 44 years for women, respectively.

Nowicki et al. (1996) found in a cadaver sample the stenotic intervertebral foramen to be most frequent at L5 / S 1, whereas the occult or resolved intervertebral foramen showed no preference within the lumbar vertebral levels. Eisenstein (1977) found in a skeletal sample a total of over 6Yo with suggested stenosis in at least one of the two main spinal canal diameters.

Major etiologies for spinal stenosis are congenital or degenerative reasons, rather than tumorous conditions or traumatic pathologies. One possibility is disc hemiation, which occurs commonly in the lower lumbar spine at the postero-lateral border of the disc and alters the intervertebral foramen size. Pallis et al. (1954) describe osteoarthritis to be the main cause of foraminal stenosis.

Neurologic symptoms may be caused either by direct nerve impingement e.g., the nerye root or by compression of adjoining vascular structures (Bailey and Casamajor,

1911; Putti, I92l; Brain, 1948; Dommisse, 1974; Rauschning, l98li Hoyland et al.,

1989; Gaskill et a1.,1991). Rydevik et al. (1984) propose an etiological model of initial trauma due to e.g., herniated disc, causing oedema and other acute and chronic effects including local ischemia, which finally leads to a dysfunction of the nerve fibres.

These etiologies have been shown in various clinical reports (Bailey and

Casamajor, 1911; Putti,1927; Mixter and Barr, 1934;Brain, 1948; Epstein et al., 1962;

Epstein et a|.,1964; Jones and Thomson, 1968; Cäc et al., 1980; Kirkaldy-WllIis et al.,

1982; Dorwart et al., 1983; Ogino et al., 1983; Weisz and Lee, 1983; Vanderlinden,

F. J. Rühli - Osteometric Variation of the Human Spine 64 19g4; An and Glover,1994; Avrahami et a\.,1994; Jeanneret and Jeanneret, 2002); since the first report of nerve root compression due to osteoarthritis and in the absence of tumorous or fracture etiology has been published (Bailey and Casamajor, 1911). This in early report already highlighted the frequent involvement of the intervertebral foramen

various such cases of spinal neural compression. Huizinga et al. (1952) label, due to the defacto relative morphological approaches trying to clearly define it, clinical stenosis to be rather a non-absolute concept. The shape and partially size of the vertebral endplate

lumbar was found to influence the prevalence of herniated intervertebral discs in the low

spinal region (Harrington et a1.,2001). Harrington et al. (2001) have linked a circularþ

acting shaped vertebral endplate, with its increased anular tension forces together with

f'orce vectors especially in large males, to such pathologies. They did not find a

correlation between individual stature, weight or body mass index and the presence of a

herniated low lumbar intervertebral disc. Harrington et al. (2001) also discuss if an

..inherited morphol0gic factor" may be involved in this etiological p:uzzle.

A possible etiological influence of the extrinsic vascular supply in the

(1948) pathogenesis of spondylotic myelopatþ was raised by Ogino et al. (1983). Brain

nature argues that the initial alterations by protruded intervertebral discs are of circulatory

most likely involving the venous system by causing an oedema. The arterial system,

implicated according to him, either would be involved indirectly at alatet stage or will be

(1944) directly by mechanical compression due to protrusion or osteoarthritis' Magnuson

as a mentions inflammation e.g., of the joint capsules and the ligamentum flavum'

the possible cause for lower back pain. Hasue et al. (1983) list intraneural fibrosis of

spinal nerve roots and ossifications of the ligamenta flava and posterior longitudinal

ligaments as further possible etiologies causing lower back pain and radicular symptoms'

65 F. J. Rühti - Osteometric Variation of the Human Spine Kikuchi et al. (1984) already studied the pathophysiology of radicular pain. They discuss,

based on clinical as well as cadaveric cases, a plethora of possible etiologies of

congenital, acquired, or both combined backgrounds. Vanderlinden (1984) reports

selected clinical cases of compression of the dorsal root ganglion causing sciatic pain.

Rauschning (1987) mentions disc bulging, altered ligamentum flawm and degeneratively

changed facet joints as main contributors in narrowed lumbar root canals, similar to the

reported lrndings by Nowicki et al. (L996). Putti (1927) highlights the mismatch in size

between the intervertebral foramen and the exiting nerye root, making especially the

lowest lumbar levels lulnerable to clinical conditions. Hoyland et at. (1989) propose the

hypothesis that venous obstruction e.g., due to a herniated disc, may cause periradicular

f,rbrosis and subsequently clinical symptoms. This is similar to an etiological multifactor

model proposed by Rydevik et al. (1984). Already Gill and White (1955) reported the

etiological connection between the presence of a transitional last lumbar vertebra and the

occurrence of lower back pain. Also MacGibbon and Farfan (1979) found a link between the presence factors such as a transitional lumbar vertebra, rudimentary ribs or size of transverse process and lower lumbar degeneration. The shape of the lumbar vertebral

endplate was found to be linked with disc hemiation (Harrington et a1.,2001). Metabolic etiologies, such as Vitamin D dehciency (Al Faraj and Al Mutairi, 2003), correlate with lower back pain as well.

Various reports already examined the possible morphologic difference between healthy and pathologic individuals with regard to lower back pain. Drinkall et al. (1984) report a significant difference of the sagittal spinal canal diameter for patients with lower back pain and control groups, with the former one having narïower values. Stockdale and

Fiday (1980) also found in their ultrasound based study differences between

F. J. Rühli - Osteometric Variation of the Human Spine 66 the latter ones having a naffower asymptomatic and symptomatic individuals with (i980a; 1980b) did not report in a oblique sagittal diameter aLL5. Larsen and Smith body diameters in lower back pain myelographic study any altered lumbar vertebral

making the involvement of spinal canal patients in comparison to a neutral control group,

independent of the main vertebral body size changes to be found in such clinical case

differences in vertebral carral lvertebral dimensions. In contrast, Ramani (1976) describes individuals in an X-ray based study' body ratio between asymptomatic and symptomatic

Porteretal.(1980)emphasiseintheirultrasoundstudythatthesizeofthespinalcanalis

neurogenic claudicatio than in classic more cruciai in cases of disc symptomatology and found a change in intervertebral root entrapment syndrome. Steph ens et al. (1991)

shape to being more of auricular and foramen size from either round or auricular in

also Figure l' teardrop shape in cases ofspinal pathologies; see the bony exit of the nerve root Foraminal stenosis is defrned as the narrowing of findings and symptoms patients may have radicular pain with or without sensori-motor

the spine (Yoo e/ al'' 1992; Inufusa e/ usually exacerbated with extension movements of

These radiculopathies are caused, al., !996;Humphreys et al',1998; Chun g et a1.,2000).

amongothers,byischemiaordirectnerverootimpingement(Ciricetal''1980;Kirkaldy- 1991). IFloyland et al' Willis et al.,l9g2; Resnick, 1985; Group and Stanton-Hicks, foramen venous plexus the (1989) link the mechanical obstruction of the intervertebral which would fìnally cause clinical subsequent ischemic related periradicular fibrosis,

syrnptoms

the influence of static and dynamic The quantitative and qualitative assessment of foramina has been reported for body positions on the dural sac and the intervertebral Epstein et al', 1964; Salamon et al'' various radiographic techniques (verbiest,1954; ö/ F. J. Rühti - Osteometric Variation of the Human Spine 1966; Jones and Thomson, 1968; Park, 1980; Kirkaldy-Willis e/ al., 1982; Yital et al.,

1983; V/eisz and Lee, 1983; Bolender et al., 1985; Liyang et a1.,1989; Nowicki et al.,

1996; Wildermuth et aL.,1998; Chung et a\.,2000; Fujiwara et a|.,2001). A correlation between the collapse of the intervertebral disc height and its possible clinical symptoms has alreadybeen shown as well (Hasegawa et al., 1995; Lu et al.,2000). Nevertheless,

Cinotti et al. (2002) doubt the alteration by a narrowing of the disc space and the intervertebral foramen width reduction. According to them, it influences mainly the height, whereas the intervertebral foramen width is mostly correlated with the sagittal diameter of the spinal canal and the pedicle length. Nowicki et al. (1996) emphasise the fact that an abnormal intervertebral disc is significantly correlated with stenotic foramen in the lumbar cadaver spine. Salamon et al. (1966) link the acute nerve root pain rather to herniated discs compromising the fossa below the nerve root than to the inflammation of the root itself.

Clinical and dynamic assessment of the spinal neural pathways

No exact characteristics exist, which mark the transition from asymptomatic to symptomatic in the spine (Wolf et al., 1956; Burrows, 1963; Postacchini et ø1., 1983;

Porter et al., 1987; Humphreys ef al., 1998). Clinical evaluations of the osseous spinal neural pathways have been reported in a plethora of studies. One of the changes to be associated with spinal stenosis seems to be inferior facet hypertrophy, with the major changes occurring in the middle of the intervertebral foramen (Humphreys et al., 1998).

According to Humphreys et al. (1998) the spinal nerye is forced, due to this inferior facet hypertrophy to the superior, more frequently, or to the inferior part of the foramen. Since these foramen areas are small, nerve compression and, consequently, clinical symptoms

F. J. Rühli - Osteometric Variation of the Human Spine 68 can occur. Kirkaldy-V/illis e/ al. (1982) stress the fact that the exiting spinal nerye root is

especially vulnerable while passing below the pedicle, while Púti (1927) emphasises the

finding that the most lower l-umbar levels are particularly susceptible to such a neural

entrapment. The particular anatomy of the cervical spine in relation to possible

pathologies involving neural pathways has already been addressed by Veleanu (1975),

whereas Crock (1981) and Bose and Balasubramaniam (1984) addressed it for the lumbar

spine.

As one example, the inter-individual variability of the transverse spinal canal

diameter varies, apparently mostly depending on vertebral level rather than age or sex

(Hinck et al., 1966). From a clinical perspective, the interpedicular distance increases in

cases of spinal tumors (Elsberg and Dyke, 1934), whereas Drinkall et al. (1984) found

smaller values of lumbar sagittal spinal canal dimensions for lower back pain sufferers than for control groups. Similar are the findings for a coal miners sample, as presented by

Macdonald et al. (1984), where smaller spinal canal diameters were correlated with higher lower back pain morbidity. Additionally, in an ultrasound study by Porter et al.

(1978a) the symptomatic individuals showed signihcantly smaller oblique sagittal spinal canal diameters than the asymptomatic ones. But, according to Drinkall et al. (1984), the sagittal spinal canal diameter cannot be used for the management or the prognostic value of lower back pain. Furthermore, Legg and Gibbs (1984) could not find a clear link between individual anthropometric characteristics such as stature and body weight and spinal canal size. Stockdale and Finlay (1930) describe in their ultrasound based study differences in symptomatic versus asymptomatic individuals especially in form of a namower sagittal spinal canal diameter at L5 in the latter group. In another large ultrasound study, Porter et ø1. (1980) found that the size of the spinal canal does not

F. J. Rühli - Osteometric Variation of the Human Spine 69 correlate with occupation, therefore, an altered spinal canal might be more likely due to

an ontogenetic rather than degenerative etiology. In general, s¡rmptomatic individuals

show more often in ultrasound imaging a narrow spinal canal than their asymptomatic

counterparts (Porter et al., 197 8a).

Furthermore, àEe, associated disc pathology, a trefoil-shape of the canal,

degenerative vertebral bars, soft tissue alterations and instability contribute as well

(Porter et al., 1978a; Porter et al., 1980). A strong correlation exists between vitamin D

def,rciency and lower back pain in areas with such endemic vitamin shortage (Al Faraj and

Al Mutairi,2003). Also Macdonald et al. (1984) found that smaller spinal canal

dimensions are linked with higher back pain morbidity. The size of the oblique sagittal

spinal canal dimension correlates with the treatment in symptomatic individuals but the

size of the L5 lumbar canal does not correlate with the intra-operative findings (Porter e/ al., 1978a). In an X-ray based study, Ramani (1976) reports differences in spinal canal I vertebral body ratios between asymptomatic and symptomatic individuals, with the latter ones having narrower spinal canals. As already mentioned above, Dommisse (1974) emphasises that the narrowest osseous spinal canal dimensions in the mid-thoracic region correlate with the region where the vascular supply for the spinal cord is the least, causing in some cases paraplegia. Eisenstein (1977) reports anuniform shape and capacity of the lumbar spinal canal, regardless of sex or inter-populational background. No significant difference, between a symptomatic and a control group, in lumbar vertebrae diameters have been reported by Larsen and Smith (1980b). On the other hand, the occurrence of the anatomical variation of the trefoil shaped lumbar spinal canal can vary between sex and inter-populational groups (Eisenstein, 1980). In general, Kikuchi et al. (1977)

F. J. Rühli - Osteometric Variation of the Human Spine 70 highlighted already the fact that the lumbar osseous spinal canal shows a high variability in size and shape.

A plethora of cut-off points for pathologic spinal neural pathways has been proposed so far (Elsberg and Dyke, 1934; Larmon, 1944; V/olf e/ al', 1956; Epstein et al',

1964; Hinck et al., 1966; V/illiams, 1975; Kikuchi et al., 1977; Portet et al', I9l8a;

Ullrich et a1.,1980; Kirkaldy-V/illis et a\.,1982; Bolender et aL.,1985; Hasegawa et al',

1995; Lee et a1.,1995; lnufusa et al., 1996). Postacchini et al. (1983) doubt the value of existing cut-off points in neural canal size. According to their findings, there is also no clear correlation between the presence of a trefoil shape and the mid-sagittal dimension of the spinal canal. The depth of the lateral recess decreases caudally and seems to be linked

to the shape of the neural canal and the pedicle length. Furthermore, according to them

the last two lumbar levels show the biggest normal variability. The interpedicular

distance is always bigger than the mid-sagittal one, making the later one, according to

postacchini et al. (1983), the clinically more vulnerable. They also describe the presence

of at least some relationship between the mid-sagittal neural canal dimensions and

interpedicular distance and vertebral body size. Additionally, Postacchini et al' (1983)

also found abnormally sized lateral recesses in cases of normal neural canal dimensions,

and the lateral recess size in an individual with ontogenetically altered neural canal

dimensions may be more easily affected in pathologic situations.

The overall high prevalence of radiologically detectable cervical spinal

pathologies has been showed by Pallis et øt. (1954). Surprisingly, after the age of 50,

neither the incidence nor the severity of canal or foraminal narrowing increased in his

sample of patients without neurological symptoms. Beside age per se, they discuss other

possible etiological factors such as spinal arteriosclerosis or fibrosis as well'

71 F. J. Rühti - Osteometríc Variation of the Human Spine proposed by weisz and Lee (1983) The concept of "spinal reserve capacity", as

a certain extent the absences of morphological for the lumbar spine, allows to correlate to pathological positional situations' In the reserve space with the ability of coping with when the spinai reserve capacity is elderiy and in cases of ,pi,,ul canal narrowing, (1983), clinical symptoms of iower back pain reduced, as proposed by Weisz and Lee to show the highest variability of the may occur. Also the lowest lumbar level seems defy the correlation of measured oSSeouS spinal canal reserve capacþ, which may 1983)' spinal cord morphometry (Weisz and Lee, diameters and, based on this, assumed

a result of lack of canal capacity or more of How far spinal stenosis as a clinical entìty is et al', lg52)' Dissimilar pattems and its neural content is stili unclear (Huizinga of patients and healtþ subjects (Dvorak er signifrcant differences can be found in motion

al.,1993). rotation change the relationship of Flexion, extension, lateral bending and axial

disc to the spinal nerve (Vital et al'' 1983; the ligamentum flavum and the intervertebral et a1.,1989; Nowickj et al,, |996; Louis, 1985; Liyan g et a1.,1989; Mayoux.Benhamou The non-pathologic spine shows a range in Schmid et al.,lggg; Fujiw ara et a1.,2001). ately 70", with the majority of it motion from flexion through extension of approxim

beinglocalisedinthelowermostspine(Park,1980).Thethicknessoftheligamentum

(Vital et a|.,1983;Nowicki et al',1996; Schmid flavum increases bilaterally in extension of et a:.,200|). Besides a described asymmetry et a..,1999; Chun g et a:,,2000; Fujiwara

et at' (1989) found a significant decrease the right and left foramen, Mayoux-Benhamou whereas the flexion position shows the of the intervertebral foramen size in extension, in lumbar intervertebral foramen size have been opposite. similar findings of altered size

to them, after a modelled intervertebral disc reported by schmid et al. (1999). According

72 Spíne F. J. Rühti - Osteometric Variation of the Human collapse, these positional differences were much smaller. Mayoux-Benhamou et al.

(1989) report no significant differences of intervertebral foramen sizes at various lumbar levels. For the cervical spine, Yoo et al. (1992) described an increase in the foramen size caudally. Yoo et al. (1992) also stress the fact that ipsilateral rotation increases the narrowing of the intervertebral foramen. This is particularly important, according to them, since, most cervical spine movements are combined multi-planar ones. Fujiwan et ø1.

(2001), Mayoux-Benhamou et al. (1989), Nowicki et al. (1996) and Yoo et al. (1992) addressed in dynamic cadaver studies the alterations of the intervertebral foramen dimensions in extreme extension, flexion and rotational pose, the major positional influences on human intervertebral foramen widths. Rauschning (1987) also examined the influence of positional movements on the lumbar intervertebral foramen in a cadaver based study. Liyang et al. (1989) showed, in a cadaver lumbar spine study, that in flexion not only the capacity of the spinal canal increased but also the length of the spinal canal and the posterior height of the intervertebral discs. Veleanu (1972; 1975) reports the impact of rotational movements on the cervical spine and how the transverse process helps to block non-physiological positions. Whereas extreme form of flexion mainly causes high tensions within the posterior ligaments, extreme lordosis has a high impact on the apophyseal joints (Adams et al., 1994). Lumbar lordosis was not found to be linked with lower back pain (Murrie et a1.,2003). Flexion of cervical (Yoo e/ al., 1992) or lumbar spine (Panjabi et al., 1983; Liyang et aL.,1989; Nowícl

Nowicki et ø1. (L996) widely addressed the effect of body positions such as flexion, extension, lateral bending and axial rotation on the lumbar intervertebral foramen

F. J. Rühli - Osteometric Variation of the Human Spine 73 dimensions, whereas Veleanu (1972; 1975) focused on the particular situation of movements in the cervical spine. Fujiwara et al. (2001) did a similar study on the impact of various body positions on the intervertebral foramen size in motion segments of the lumbar cadaver spine. In case of degenerative alterations, Panjabi et al. (1983) found a higher decrease in size in physiologic-dynamic situations, such as rotational movements, for intervertebral width than height.

The bulging of the intervertebral disc is, according to Reuber cited by Panlabi et al. (1983), in case of degenerative lumbar spine approximately 2 mm. An artifìcial collapse of the intervertebral disc, which anatomically influences the neural pathways less than a degenerative disc with subsequent fattening and protrusion, decreased the relative changes in relation to the two extreme positions (Mayoux-Benhamou et a1.,1989).

Computer assisted simulation of narrowing of intervertebral disc space to determine the relationship between intervertebral disc height, which is greatest anteriorly, and the size of the intervertebral foramina, with a I mm narrowing leads to a reduction of

20Yo to 30Yo of the foraminal area (Lt et a1.,2000).In a similar study by Cinotti et al.

(2002), the artificial narrowing of the disc space caused mainly a decrease in intervertebral foramen height, rather than foramen width. The latter one was more linked to the sagittal diameter of the spinal canal or pedicle length.

The use of a chronic compression model in rats allowed Iwamoto et al. (1995) to

explore the sequence of pathologic alterations in lower lumbar spinal compression.

Surprisingly, at the very beginning of such a process, the epidural blood vessels are

damaged and only later in the long-lasting process the nerve roots are injured.

Cross-sectional areas of the non-pathologic spinal cord in cadavers have been published earlier (Lassek and Rasmussen, 1938; Elliott, 1945; Kameyama et al., 1992).

F. J. Rühli - Osteometric Variation of the Human Spine 74 Scoles et al. (1988) provide gold-standard data of the non-pathologic spinal canal dimensions in the macerated thoraco-lumbar spine. Bolender et al. (1985) provided radiological values of cross-sectional areas of the dural sac and spinal canal diameters in symptomatic patients. Inufusa et al. (1996) report a significant correlation between the mid-sagittal diameter of the spinal canal and its cross-sectional area on the lumbar level.

Inufusa et al. (1996) report as normal value for the lumbar spinal canal an overall cross- sectional area of 200 mm'. The measuring the mid-sagittal diameter, as done in their study, allows estimating overall canal size. These sizes show for the neural tissue no significant correlation with body weight but with body height, and a remarkable inter- individual variation. The relative cervical cross-sectional area, according to Kameyama e/ al. (1992), is alike within individuals. According to Gepstein et al. (1991) the sagittal diameter of the spinal canal is the only parameter of a series of osseous vertebral dimensions, which correlates with the cross-sectional area of the spinal canal. Eisenstein

(1971) recommends focusing on the absolute values in sagittal canal dimensions, which are more crucial than the transverse diameter or any ratios with the vertebral bodies. This view of mainly the sagittal diameter of the spinal canal being the clinically critical measurement, is also supported, at least for the cervical spine, by Wolf et al. (1956) or for the lumbar spine by Kikuchi et al. (L977).

A clear relationship between intervertebral foramen height and sagittal diameter of the spinal canal was reported by Epstein et al. (1964), which they declare to be a crucial factor, together with a tendency of narrowing in the lateral recess, in the occurrence of clinically relevant spinal diseases. In an earlier report, Epstein et al. (1962) discuss the importance of decreased lateral spinal canal recess size in the occurence of clinical lower back symptoms. Such a variation in lateral recess size can be found, according to them in

F. J. Rühtí - Osteometric Variation of the Human Spine 75 approximately I}-ISYo of all individuals. The particular shape of the lateral recess has also been addressed widely by Kikuchi et al. (1977). Eisenstein (L977) summarizes in his lumbar spine study that the osseous narowing of the spinal canal, as the only reason for spinal stenosis, may not be correct. Spinal stenosis will affect more the intervertebral foramen than the main vertebral canal. ln another study by Eisenstein (1980), he rules out facet osteophles or trefoil configuration of the lumbar spinal canal as main etiologies for nerve root compression.

The clinically crucial and unique patho-anatomical features of the intervertebral foramen have already been addressed by Magnuson (1944) and Rauschning (1987).

Surprisingly, Magnuson (1944) found the root ganglion in fresh cadavers to filI out the vast majority of the foramen and, furthernore, he describes a high variability of the anatomy of the intervertebral foramen and its content. Hasegawa et al. (1995) found, in their cadaver study of the lumbar spine, a significant correlation between posterior intervertebral disc height and foramen height. Furthermore, they found a correlation between foraminal cross-sectional area and the nerve roots size. The ratio of these two measurements was higher, according to Hasegawa et al. (1995), in individuals with a possible nerve root compression. Additionally, in the possibly affected subgroup the posterior disc height as well as the foraminal height was generally smaller. As Putti

(1927) already stated, there seems to be a mismatch between intervertebral foramen space and spinal nerye size particularly in the two lowest lumbar segments. Ebraheim et al.

(1996) provided cadaver and macerated cervical intervertebral foramen dimensions acquired in neutral position, which can be used as reference data. In a cadaver study,

Hoyland et al. (1989) suggest that mechanical occlusion of the intervertebral foramen venous plexus could lead via ischemia to periradicular f,rbrosis and, therefore, to clinical

F. J. Rühli - Osteometric Variation of the Human Spine 76

better school test scoring (Porter et a1.,1987). Furthermore, workers with smaller spinal

canal dimensions show in general a higher lower back morbidity (Macdonald et al.,

1e84).

Historical perspectives of spinal disorders and morphometry

A historic perspective on the spinal morphology is still not widely used. In a

archaeologic pilot study exploring the possible influences of individual hardship during

growth on juvenile spinal canal dimension, Porter and Pavitt (1987) describe several

significant links between individual skeletal or dental stress markers such as Harris lines,

cribra orbitalia, porotic hyperostosis or dental hypoplasia. Noteworthy, they found a

positive correlation between the decrease of mid-sagittal spinal canal size, which is the

most important clinical diameter of spinal neural pathways, and the occurrence of Harris

lines on most lumbar levels (Porter and Pavitt, 1987). Porter and Pavitt (1987) postulate

that unknown factors acting on the foetal development of the individual spinal canal may

also result in a susceptible immune system. Therefore, the latter could explain the link

with the occturence of Harris lines, since Harris lines are in general to be more frequently

found in cases of severe acute infection or poor diet. The secular change ofneural spinal

pathways in a cultural transition period from a hunter-gatherer to a settled agricultural

society in North America was examined by Clark et al. (1985). They found a slightly

smaller sagittal spinal canal dimension in the thoracic and lumbar spine in the agricultural

society, even after controlling for sex and age. For the transverse diameters, only females had smaller dimensions, \Mhereas males had higher values than their hunter-gatherer counterparts. The agricultural males also had larger lumbar vertebral body heights; which was less expressed in the thoracic spine and with an opposite trend for females (Clark et

F. J. Rühli - Osteometríc Variation of the Human Spine 7g al., 1985). Clark et al. (1985) state that the lumbar sagittal spinal canal dimension is an excellent indicator for disrupted growth in individual's early life. This dimension is not correlated with tibial length, whereas transverse diameters are. Both, tibial length and vertebral body height did, according to the results by Clark et al. (1985) not change during a cultural shift. Since the thoracic spine completes more of its growth in the prenatal stage than the lumbar spine, it is not surprising that the first one shows stronger correlations between main spinal canal dimensions and vertebral body height (CIark et a1.,..1985). Clark et al. (1985) describe more conelations of various osteometric spinal assessments. Sagittal and transverse spinal canal diameters are correlated as high as transverse spinal canal diameter and age groups. Furthermore, transverse spinal canal diameter is correlated with sex, unlike the sagittal diameter. Posterior vertebral body height is correlated with sex and cultural transition. Finally, anterior vertebral body height is also correlated with cultural change and with posterior vertebral body height. Clark et al. (1985) also found that intervertebral foramen width is only correlated with sagittal diameters of the spinal canal but not with the transverse diameter of the spinal canal or with vertebral body height. These correlations seem not to be clouded by variables such as sex, age or culture (Clark et al.,1985). The shift from a protein-rich hunter-gatherer society to a protein-poorer agricultural life style, as examined by Clark et al. (1985), results in smaller spinal canals. This is more strongly expressed in the sagittal dimension, which is more vulnerable to influences in the pre-and neonatal growth period and more visible in the lumbar spine (Clark et a|.,1985).

In an osteometric studies including two Early Medieval samples from present

Poland, Piontek (1973), found a strong correlation for all vertebral levels between sagittal and transverse diameters, but no such significant relationship exists for the majority of all

F. J. Rühli- Osleometric Variation of the Human Spine 79 vertebral levels between these two diameters measurements and the vertebral body height.

Piontek (1973) describes, with just a few exceptions, a correlation between the transverse vertebral body diameter and the transverse spinal canal diameter. This seems, according to him, not to be true for the sagittal dimensions of these two structures. Another study briefly focusing on historic spinal morphometry is the one by Tatarek (2001).

From a historic perspective, changes in the prevalence of degenerative spinal diseases have been linked to possible alterations in cultural and, therefore, mechanical loads (Larsen, 1980; Larsen, 1981; Larsen, 1982; Bridges, 1991). Larsen (1980; 1981;

1982) mentions a signihcant decrease of cervical and lumbar degenerative joint diseases, with a reduction of up to 27Yo of its prevalence, from a pre-agricultural hunter-gatherer society to a settled corn dependent agricultural community, both located in the same

American costal area. He explained this as being related to a decrease in mechanical stress due to the change in life-style (Larsen, 1980; Larsen,1982).

The osteometric definitions of spinal landmarks allow comparison of data with various geographic and historic backgrounds (Aeby, 1879; Anderson, 1883; Rosenberg,

1899; Wetzel, 1910; Hasebe, 1913; Thomson, 1913; Cy.ia*, 1920; Stefko, 1926; Jacobi,

1927; Mafün, 1928; Frey, 1929; Matiegka, 1938; Wood-Jones, 1938; Lanier, 1939;

Huizinga et a\.,1952;Fruncis, 1955; Davis, 1961; SchuItz,196l; Stewart, 1962; Epstein

et a1.,1964; Kaliszewska, 1966; Hurxthal, 1968; Piontek and Bud4mska, 1972; Piontek

and Zaborowski, 1973; Dommisse, 1974; Dommisse, 1975; Heim, 1976; Cwirko-

Godycki and Swedborg, 1977; Eisenstein, 1977; Kikuchi et al., 1977; Riegerova, 1979;

Tibbetts, 1981; Postacchini et aL.,1983; Nissan and Gilad, 1984; Amonoo-Kuofi, 1985;

Trinkaus, i985; Cottenll et al., 1986; Gilad and Nissan, 1986;Nakashima, 1986; Nissan

and Gilad, 1986; Berry et ø1.,1987; Porter and Pavitt, 1987; Minne et al., 1988; Scoles e/

F. J. Rühli - Osteometric Variation of the Human Spine 80 et 1995; al., 1988; Gepstein et al., 1991; Sanders, 1991; Jankauskas, 1994;Lee al',

Tominaga et a1.,1995; Xu et al.,1995; Sanders, 1998; Tatarek, 2001).

there is not Nevertheless, as Katz et at. (1975) stated at least for the cervical spine,

(1988) this fact in much data available e.g., on u.n"b.ul body size. Scoles et al. highlight

its terms of the absence of knowledge on the thoraco-lumbar spinal morphometry, despite

crucial need of it e.g., in orthopaedic surgery'

This lack of morphometric information is striking especially if one is aware of the

disorders linked with importance, such as in modern clinical medicine, of human spinal on spinal short-term morphologic mal-adaptations. Furthermore, this lack of knowledge

neural evolution is surprising in particular for the macerated intervertebral foramen and

one can rely canal dimensions. At least for the cervical intervertebral foramen dimensions

standard on data published by Ebraheim et al. (1996). Nevertheless, the well-established

for the measurement schemes by Hasebe (1913) and Martin (192s) provided definitions

measurement of the spinal canal diameters only'

to possible Surprisingly, no study including historic specimens paid full attention

of the intervertebral secular trends in spinal neural pathways dimensions. The assessment pathophysiology of foramen is crucial as its alterations play a significant role in the

radiculopatþ or spinal stenosis, main etiologies of back pain, which, cause enoÍnous 2000). No study costs in industrialized countries health care (Maniadakis and Gray, post- exploring a possible secular alteration of the intervertebral foramen in

the superior industrialization societies exists. since the inverted teardrop-like shape of

(swanberg, 1915; Panjabi and inferior soft tissue parts of the intervertebral foramen space different from et a1.,1983; Vital et a1.,1983; Rauschning, 1987; Inufusaet al',1996) is

(ciric et 1980; Mayoux- its osseous outline, earlier proposed clinical measurements al.,

81 F. J. Rühti - Osteometric Variation of lhe Human Spine between the dental hypoplasia in an individual and a small lumbar interpedicular distance or between the presence of a small sagittal diameter of the spinal canal and the prevalence of Harris line.

The non-human spine

Various functional and morphological aspects of the non-human spinal column have already been addressed (Keith,1902; Wetzel, 1910; Nathan et al., 1964; Mehler,

1969;Fartan, 1978; Cotterili et a\.,1986; Fox and Wilczynski, 1986; Pun e/ al., 1987;

Shapiro, 1993; Maclarnon, 1995; Shapiro, 1995; Tominaga et al., 1995; Maclarnon,

I996a; Maclarnon, 1996b; Sanders, 1998; Boszczyk et aL.,2001; Kandziora et a1.,2001;

Argot, 2003). Animal spines have been used as models for the human spine for various reasons. Both, the cervical spine of sheep (Kandziora et a1.,2001) as well as the one of the baboons, at least as highlighted by Tominaga et al. (1995) show to a certain degree similarities to the human spinal anatomy.

The increase in spinal cord size during primate evolution is explained by

Maclarnon (1996a) most likely due to increased complexity in locomotion. Both, Homo sapiens and Pan troglodytes show a sudden end of the spinal cord, most likely due to the absence of any tail. The expansion of the corticospinal tract, only to be found in mammals and important for fast and smooth activities (Towe, 1973), and of the dorsal columns, consisting of afferent sensory neryes, within the spinal cord, could be the reasons for the increase in cervical and thoracic spinal cord dimensions during primate evolution

(Maclarnon, 1996a). The human lumbar spinal canal shows, according to Maclarnon

(1995), even more particularities, such as the lack of any decrease in diameter towards its caudal end. Maclarnon (1995) explains this as being a result of intrinsic and / or extrinsic

F. J. Rühti - Osteometric Variation of the Human Spine 83 influences, such as bipedialism, acting on the vertebral canal. The osseous human spinal canal, therefore, does not reflect its neural content as it does in other primates. This is only true for the lumbar segment, since for the more cranial parts such a correlation

apparentiy does exist (Maclarnon, 1995).

Surprisingly, in comparison with other primates humans tend to have large and wide pedicles in relation to their pedicle length and body size (Shapiro, 1993). At least

this distinctive human pedicle morphology may be resulting from the unique pattern of

locomotion (Shapiro, 1993). Furthermore, human lumbar pedicle morphology may echo

bending forces and may by influenced by the presence of the ilio-lumbar ligament (Davis,

1961; Shapiro, 1993).

The particular spinal anatomy with its bulþ lower back muscles, the functional

lordosis and a more dorsal displacement of the posterior spinal ligaments, makes humans

able to handle much higher weight bearing than their primate relatives (Farfan, 1978).

As a side issue of this work, which will not be further addressed, CT scans of

some selected ape cadavers have been performed to illustrate the in vivo spinal

morphology, and in particular the relation between vertebral body height and

intervertebral disc dimensions in the lumbar spin; see also Figure 2.

F. J. Rühli - Osteometric Variation of the Human Spine 84 (:Èr Ifft' cH il'l['. & Ptnrs. 2 i tr¡ ì krìou¡ t, 17 na\J ?OO? : rrì k rìourì - 13:43:23 CT

R L

kvP: ::ìOO 'rA rrrgec : !-tOO [ìfrs : l50 i t rc¡O lhk:1 n¡¡¡ ?oo/500 IHJ a) Chimpanzee (Pan troglodytes)

oRÉìN(; Ht-5t t Pl rìrq 1 )utì - 1 1 l'1,:rr1 13:24:73

R d;

kVF:12O uA: iOO Dlljcc :r-ìoaJ ilrfì.j : l'-)O Vilr.:.¡3 lltk:2 r¡r¡tt U/l i l?OOi'.)OO

b) Orangut an (P ongo py gmaeus)

Figure 2 CT based 3-D-surface reconstructions of thoracic and lurrbar spines of selected ape cadavers

F. J. Rähli - Osteometric Variation of the Human Spine 85 (ìilìBON (;ilìrtON t)r Jor res & [' t.rì.s . :J ReF:rr¡krroqtr- 11 lltrg 2OO2 Rlrcl : rrr r kr routr. 1.1:l)?:l)l CT

Æ.' R &;

kVP: t20 r¡A: J¡)O rr¡u,:t-: t>OO rrl^¡s:150 V iLr c¡¡0 I l¡k: 1 ¡rrr H/l:12OOll>OO ñi c) Gibbon (Hylobates, species unhtown)

Figure 2 (cont.) CT based 3-D-surface reconstructions of thoracic and lumbar spines of selected ape cadavers

F. J. Rühli - Osteometric Variation of the Human Spine 86 Earlier reports (Keith, 1902; Schultz, 196I) addressed the variation in the non- human spine in comparison with its situation in humans' Bohart (1929) already mentioned the fact, that sacralisation of the lower lumbar spine is not only very frequent -often primate in humans, but even more to be found in some species of monkeys. The

body spinal cord does not differ from the one in other mammals with regard to its size / weight ratio, also there is just a small variation in relative cord length in primates

(Maclarnon, 1996b). Primate spinal cord weight and length are strongly correlated with body weight (Maclarnon, 1996b). Shapiro (1993) examined the features of the vertebral body surface areas and pedicle dimensions among primates including an Australopithecus africanus individuai and anatomically modern human samples. The influence of unique human posture and locomotion was in general found to be weaker than expected

(Shapiro, 1993). To summarize,Farfan(1978) concludes that the human spine is from an

evolutionary perspective a well adapted structure'

Major evolution of the human spine and its physiological adaptations

The human vertebral column has evolved from the ones of other primates by

adaptations possibly linked with changes in life-style and environmental habitat'

Boszczyk et at. (2001) highlights the fact that humans, in comparison to their closest

living relative, the chimpanzee, show a functional adaptation to the higher axial loading,

mostly by ar increase in the transverse rather than the sagittal vertebral body diameter'

This allows humans to have a relatively large surfac e atea, especially in the lumbar spine'

Mehler (1969) mentions not only the increase of the spino-thalamic tract during mammal

evolution, but also as a cut-off between the neuronal tract of humans and chimpanzee

87 F. J. Rühti - Osteometric Variation of the Human Spine versus other primate and non-primate spines, the absence of the spino-olivary connections in the formers, with this fact possibly being a reflection of bipedal locomotion.

Another striking evidence of unique human neuronal evolution has been reported as the loss of a sialic acid'(Varki, 2001). Other possible examples of biochemical evolution in humans (Rühli and Henneb erg, 2001; Rühli and Henneberg, 2002) will be addressed later in this work.

The particularities of the human lumbar spine in relation to it closest relatives have been highlighted by Farfan (1978), by emphasizing the great functional variability

human due to e.g., greater thickness of lumbar discs. Schultz (1961) pointed out, that the

spine shows, due to the particular posture and its related mechanical implications, very

broad lumbar vertebrae. The reduction of the nuchal musculature, according to Schultz

(1961), results in exceptionally short cervical spinous processes in humans'

The spinal morphology is reflective of the amount or direction of physical forces

acting on vertebrae (Davis, 196l; Putz, 1981; Louis, 1985). In contrast to terrestrial

quadrupedal animals, the human spine is exposed to the demands of bending. The impact

of locomotion patterns such as bipedialism, posture influences and other functional

aspects e.g., loading / forces in lifting, of the vertebral column and its linked muscles

have been outlined earlier (Davis, 1961; Nathan et al., 1964;Putz, 1981; Yettram and

Jackman, 1982; Louis, 1985; Pun et al., 1987; Sanders, I99l; Putz and Müller-Gerbl,

1996; Sanders, 1998; Boszczyk et a1.,2001). It is well known that bipedialism directly

influences the arrangement of central nervous system structures such as e.g', the position

of the spinal cord in relation to the brain and thus the placement of the foramen magnum

(Schaefer, lggg). The particular interaction of physiological and pathological mechanics

and spinal anatomy has also been discussed in earlier reports (White and Hirsch, I97I;

88 F. J. Rühli - Osteometric Variation of lhe Human Spine 1982; Panjabi et a1.,1976; Farfan,l978;Nachemson et a1.,1979;Yetttam and Jackman, panjabi et a\.,1983; Louis, 1985; Silva e/ al., 1997; Sanders, 1998).

The early hominids, the Altstralopithecines. show different spinal morphology

functional than modem Homo sapiens (cook et al., 1983; Sanders, 1998), with some implications such as proposed greater massiveness of their back musculature' I osteoarthritic changes, as seen e.g., in the skeleton of the La chapelle-aux-Saints

the morphologic remains of Homo sapiens neandertalensls (Trinkaus, 1985), influence

a frequent pathology characteristics of the human vertebral column. Another example of

Morbus interfering with normal spinal architecture is juvenile kyphosis, so called

Australopithecus scheuermann, which e.g., was supposed to be present in the Al-288

africanus, afarensis skeleton (Cook et a\.,1983). Individual STS 14, anAustralopithecus

by having small showed distinctive morphological features compared to modern humans a possibly vertebral surface areas and relatively short pedicles at L6, both suggesting

small size unique locomotive pattern or being simply an allometric trait related to

canal for (Shapiro, 1993). Sanders (1991) studied the cross-sectional areas ofthe neural

decreases in size relative each level in the lumbar spine. Among hominoids, lumbar canal

possess much to centrum areas with increasing body weight. Modern humans generally

The lumbar vertebrae larger neural canal areas relative to body size than their ancestors.

body sizes of Australopithecines show smaller centra than predicted for their estimated

intervertebral foramina and relatively wide neural arches and canals (Sanders, 1991). The

with modern of the STS 14 individual are supposed to be relatively large in comparison

nerye size of STS humans, unlike its relatively short pedicles (Shapiro, 1993). The spinal

had occupied 14 may have been increased or, more likely, the spinal nerves may

humans' relatively less space of the intervertebral foramen as in anatomically modern

89 F. J. Rùhlí - Osteometric Variation of the Human Spine making a symptomatic nerve injury less likely in STS 14 (Shapiro, 1993)' The Early

Upper paleolithic individuals from Predmosti, which are of the Cro-Magnon t)'pe, show in comparison to modern samples relatively small neural pathways (Matiegka, 1938).

Stewart (1962) found no evidence for an anatomical essentially different cervical spine in

Neandertals in comparison to modem humans and describes the long spinous process at

C5 as one characteristics of the Neandertal spine. Apparently, the lower cervical spinous process became less robust. Additionally, European Neandertals show relatively shorter upper and lower limbs, as pointed out by their brachial and crural indices (Trinkaus,

1981; Ruff, 1994; Holli day, 1996; Holliday, 1997; Hotliday, 1999). It is also still debated how the scapula morphology of the Neandertals changed towards modern humans

(Churchill, 1996). Trinkaus (1985) and Heim (1976) emphasize the high robusticity of the

Neandertal spine in comparison to the one of anatomically modern humans, but both also

stress that conclusions drawn shortly after the discovery of these skeletons about its

particularities, special morphology are not correct. Heim (1976) mentions, among other

the big cervical neural canal of the La Ferrassie I Neandertals individual as well as its

robust cervical neural arch.

Some altered spinal features are expressed in modern humans e.g., as variation of

process or neural the number of vertebrae, mostly thoracic and lumbar, increased spinous

nerves in canal size, changed intervertebral disc height, changed numbers of segmental

comparison with total number of vertebrae, variation of the foramen of the transverse

process, or different proportions of the major spine regions (Keith, 1902; Horwitz,1939;

Francis, 1955; Gill and white, 1955; Bornstein and Peterson, 1966; MacGibbon and

Farfan, 1979; Cotlenll et a\.,1986).

90 F. J. Rühti - Osteometric Variation of the Human Spine

spine (Hasebe, 1913). Anatomical alterations of the same structure but on dissimilar

levels can be a result of different etiologies. Cervical lordosis, for example, is apparently

an exclusive effect of intervertebral discs, whereas the lumbar lordosis is a result of both,

the arrangement of the vertebral bodies and of the intervertebral discs (Jankauskas, 1994).

Besides obvious osseous adaptations, the non-osseous parts of the human spine

such as the ligamentous elements are expressing evolutionary adjustments as well

(Farfan, 1978). The ligamentous apparatus of the vertebral column, which includes beside

the major anterior and posterior intervertebral ligaments the annulus hbrosus of the

intervertebral disc, is of particular evolutionary relevance (Farfan, 1978).

The mechanical load bearing function of the human spine, as it is evolved into its

current physiological form, is vital. The main function of the human intervertebral discs

is to distribute equally any mechanical loading regardless of the vertebral position. In

addition, the deep back muscles as well as various ligaments support this function. The

anatomical adaptation of the spine in general can be seen as the best solution of very

competitive needs, this is stability and mobility (Darwin, 1859; Davis, 1961; Veleanu,

1972;Yeleanu,l975; Putz, 1981; Louis, L985;Putz and Müller-Gerbl, 1996; Boszczyket

al., 200I). The still ongoing evolution of the human vertebral column can also be

investigated by exploring the frequency and extent of anatomical variations and by the

occurrence and type of pathologic mal-adaptations.

Anatomical variations of the spine

Numerous variations in the occurrence, the arrangement and the function of soft tissue body parts such as muscles, vessels or visceral organs exist. Anatomical variations in the human vertebral column are rather frequent. Many studies have been conducted to

F. J. Rühli - Osteometric Variation of the Human Spine 92 explore the form and- intensity of expression of spinal variations (Rosenberg, 1899;

Dwight, i901; Keith,1902; Cyriax, 1920; Willis,1923; Willis, 1924;Putti,1927;Martrn, 'Willis, 1928; Bohart,1929; Cushway and Maier, 1929;Frey,7929; 1929; Giles,l93l;

Philipp, 1932; Stewart, 1932;'Junghanns, 1933; Horwitz, 1939; Lanier, 1939;Larmon,

1944; Magnuson, 1944; Nlbrook, 1955; Francis, 1955; Gill and White, 1955; Schultz,

1961; Epstein et ø1., 1962; Burrows, 1963; Epstein et al., 1964; Post, 1966; Salamon e/ al.,1966; Veleanu, 1975; Arnoldi et a1.,1976; Eisenstein,1977; MacGibbon and Farfan,

1979; Riegerova, 1979; Eisenstein, 1980; Susa and Varga, 1981; Tibbetts, 1981; Hasue e/ aL.,1983; Kikuchi et a|.,1984;Larsen, 1985; Parke et a|.,1994; Hoshovski, 1996; Tribus and Belanger, 2001). Willis (1929) differentiates between phylogenetic, (e.g., partial sacralisation of the last lumbar vertebra), developmental (e.g., defective spinous process) and acquired spinal variations (e.g., trauma related conditions).

The various patterns of variability of the human vertebral column can be shown among others by the variation in the number of vertebrae, the configuration of processes of the neural canal, the disposition and asymmetry of zygoapophyseal articular facets, the sacralisation and lumbalisation of the sacro-lumbar junction, the variation of the transverse foramen, the extent of vertebral fusions, the vertebral body, pedicle, spinal canal, spinal nerve or dural sac morphology, the variations of the nerve root sizes and the occuffence of additional ribs (Rosenberg, 1899; Dwight, 1901; Keith, 1902; Hasebe,

1913; Cyriax,1920; Willis, 1923; V/illis,1924;Putlri,1927; Martin, 1928; Bohart,1929;

Cushway and Maier, 1929; Frcy, 1929; Willis, 1929; Gi\es, 1931; Blumensaat and

Clasing, 1932; Philipp, 1932; Stewart, 1932; Horwltz, 1939; Lanier, 1939; Allbrook,

1955; Francis, 1955; Gill and White, 1955; Schultz,196I; Epstein et a|.,1962; Burrows,

1963; Bornstein and Peterson, 1966; Salamon et al., 1966; Veleanu, 1975; Saillant, 1976;

F. J. Rühli - Osteometric Variation of the Human Spine 93 Kikuchi et al., 1977:'MacGibbon and Farfan,1979; Riegerova, 1979; Susa and Varga,

1981; Tibbetts, 1981; Hasue et al., 1983; Postacchini et al., 1983; Kikuchi et al., 1984;

Larsen, 1985; Hoshovski, 1996).

Anatomical variation, extremely conìmon even in asymptomatic "- be individuals. Cushway and Maier (1929) found in an X-ray sample of 931 healthy men a total of 414 cases showing any osseous spinal variations. A similar percentage of approximately 45Yo of symptomless individuals expressing some sort of spinal variation was reported by Bohart (1929). Giles (1931) reports in an X-ray based study a prevalence of approximately l4To of vertebral anomalies of any form. This includes alterations of vertebral segmentation, hemivertebra, spina bifida, or the occurrence of cervical or

lumbar ribs. A high frequency of numerical vertebral variations of approximately l5o/o was reported by Allbrook (1955) for a modern East African sample, whereas Bornstein

and Peterson (1966) detected an overall variance of lIo/o, Stewart (1932) one of l2o/o,

Tibbetts (1981) for males a total of 8o/o and one of 10% for females, Willis (1923) forthe

thoraco-lumbar spine of Whites one of approximately 5Yo, Blumensaat and Clasing

(1932) in a clinical sample a total of 5Yo, Martin and Saller (1957) a total of 8o/o and Keith

(1902) mentions the same percentage. Schultz (1961) and Frey (1929), both found a total

of 3f/o and 32Yo, respectively, of any vertebral numerical variation. Dommisse (1974)

describes in a sample of six cadavers one with an additional lumbar vertebra. MacGibbon

and Farfan (1919) found in their large sample a total of 8o/o with transitional vertebra,

whereas in another osteometric study, briefly mentioned in a more clinically orientated

report by Gill and White (1955), ll% of skeletons show transitional vertebrae. Philipp

(1932) reports more than 25Yo ofa sample of pelvis specimens to have some sort of sacral

anomalies. Epstein et al. (1962) estimate l0% - 15% of all individuals showing a

F. J, Rühti - Osteometric Variation of the Human Spine 94 decreased size in the lumbar spinal canal recess size. Cyriax (1920) describes in his sample of cadaveric and macerated spines a high degree of variation especially in the vertebral size ratios. Also a high number of uni- / or bilateral alterations in numbers of vertebrae has been reported eailier by Horwitz (1939). Furthermore, he found a link in the occuffence of vertebral segmentation alterations and anatomical variations of the lumbo- sacral nerve plexus.

By addressing the blood supply of the spinal cord, Dommisse (1974), emphasizes the fact that the vascular supply shows a striking anatomical variability. Patke et al'

(1994) describe a higher variability of the arterial supply for the three lowest lumbar

intervertebral foramina than show the more cranial or caudal ones. On the other hand,

Tribus and Belanger (2001) did not frnd a variation in the occurrence but only in the

localization of the median sagittal arlery.Larsen (1985) investigated not only the

expression but also the variability in the posterior vertebral body anatomy by focusing

e.g., on the foraminae caused by the basivertebral veins and the scalloping of the lumbar

vertebrae.

A high variation in spinal nerve and intervertebral foramen arrangement was

described by Magnuson (i944) based on a sample of ten fresh cadavers. Similar reports

are provided by Hasue et al. (1983) and Kikuchi et al. (1984) stating that congenital

variations of the nerve root, such as branching or root merges, are quite common, with a

prevalence of approxim ately 9%o. Vanderlinden (1984) describes a few clinical cases with

part of the a variant location of the dorsal root ganglion, in the proximal instead of lateral

intervertebral foramen, linked to sciatic pain. Francis (1955) describes a high degree of

variation of the foramen of the transverse process. Horwitz (1939) mentions the variation

of the lumbo-sacral and posterior sacral nerve plexus and its relation to the alteration in

95 F. J. Rühli - Osteometric Variation of the Hurnan Spine number of vertebrae. Furthermore, Dunn (1912) reports a high variability for the size of cervical nerve roots in albino rats.

The trefoil shaped lumbar spinal canal, another frequent spinal morphology variant, is addressed by Eisenstein (1977; 1980) as being just an anatomical developmental modihcation, which is most frequent at L5 and more often to be found in females and in certain inter-populational group, and which is not primarily linked with local nerve entrapment and its subsequent symptoms. It is present in approximately l5o/o of all individuals at L5 (Eisenstein, 1977; Eisenstein, 1980). Postacchini et al. (1983) describe this particular shape in l6Yo of an Italian sample and in I2%o of an Indian sample, but they do not describe a correlation between the trefoil shape and the mid- sagittal neural canal dimension. Furtherrnore, Kikuchi et al. (1977) stress the fact that the osseous spinal canal shows a wide variation in size and shape. The variation of the spinal dural sac has been shown by Salamon et al. (1966), who fund its termination at Sl / 52 in

87Yo of their sample only, with its ending in other cases even further caudal.

Additionally, in forensic situations the variability of the human spinal morphology in individuals has been used for identification purposes (Riepert et al., 1995).

Summarizing, it is difficult to define a clear division in the human spine between a pathologic finding and an anatomical variation Q,{iedner, 1932; Allbrook, 1955). Bohart

(1929) did not find any correlation between the presence of any spinal variation and the likelihood of work-related back injuries. Even anatomical variations itself can be tricky to be identified, especially on X-rays (Cushway and Maier, 1929). While Giles (1931) denies a clear link between the occurence of spinal abnormalities and backaches in a particular individual, spinal variations such as transitional lumbar vertebra and rudimentary rib have been linked by MacGibbon and Farfan (1979) to low lumbar

F. J. Rühli - Osteometric Variation of the Human Spine 96 (1955) also mention a degeneration and its subsequent clinical impact. Gill and white

pain. They describe a correlation between transitional last iumbar vertebra and lower back

last lumbar vertebra. smaller sãgittal spinal canal dimension in cases of a transitional presence of Already Philipp (1932) linked the occurrence of sacral pain and the

and and white (1955) sacralisation of the rower lumbar spine. wiltis o92a; 1-929) Gill

of low lumbar vertebral alreadyhighlighted the importance of a link between the presence

to recapitulate some anomalies and the occurrence of back pain. Therefore, it is essential

of the major spinal pathologies, which may have at least a partial evolutionary

spinal mo¡phology' background and may help to better define the true normal range of

Microevolutionary and secular trends in human anatomy

The term ,.secular trend" is linguistically derived from the Latin word saeculum

changes especialiy of meaning a generation. Therefore, secular trends describe short-term

morphological traits. their active Humans are in evolutionary terms actors, which do not fully reflect

process evolution as participation (Henneberg,1997). Henneberg (1997) describes the of

society and the a feedback regulated by interactions between environment, technology'

human body. He also states, that, since our environment is self-changing and'

technology as well' additionally, influenced by us too, our anatomy may be adapted to body and its For him society and technology are acting as sieves between the human of workload or sports environment. Particularly, modern lifestyle with its unique aspects

underestimated. activities does have an influence. Its medical signihcance is repeatedly fall Morphologic body changes occurring in the modern Homo sapiens may

within various etiological categories such as anagenetic or cladogenetic microevolution

97 F. J. Rühli - Osteometríc Variation of the Human Spine (Wiercinski, l9l9). Nèw selective forces, mutagenic agents, genetic intermixtures and environmental conditions act differently on the human body.

Various influences like variation of selective pressures, exchange of genes, environment e. g., climate as in the case of the altered prevalence rate of the lateral internal thoracic artery (Surtees et al., 1989a', Surtees et al., 1989b; Henneberg , 1992), or change of socio-economical structures, such as from hunter-gathering societies to more settled communities o.g., in the Late Paleolithic-Mesolithic transition period in Central

Europe, have an impact on human anatomy, metabolism and behaviour.

Especially, gracilisation of the human body, a structural reduction of its size and bony robusticity, has been shown since the Late Paleolithic in European samples

(Schwidetzky, 1962; Schwidetzþ, 1967; Schwidetzky, 1969; Schwidetzþ, 1972;

Schwidetzþ and Rösing,1976; Vallois and de Félice, 1977;Frayer, 1980; Frayer, 1981;

Wurm, 1982; Frcyerr 1984; Schwidetzky and Rösing, 1984; Jacobs, 1985a; Jacobs,

1985b; Schwidetzþ and Rösing, 1989; Ruff e/ al., 1993; Mathers and Hennebetg, 1996;

Ruff ¿/ a1.,1997; Trinkaus, 1997). The advantage, in terms of energetic fitness, of having

more gracile bodies has already been highlighted as a possible underlying factor (Frayer,

1981; Frayer, 1984; Henneberg and Steyn, 1995). Wurm (1982) describes a decrease in

stature in historic times based on the assumption of etiologically related decreased animal

protein intake. Contrary, Larsen (1981) doubts for a historic American sample the

primary role of altered protein intake in causing a decrease of postcranial size and

robusticity, blaming diminished mechanical load to be more likely responsible. This

negative secular trend, as found in Europe, is only reversed since the early 20'o century by

a positive temporal trend in increased stature only in the Northern Hemisphere of still

debated etiology. As one of the few exceptions, no secular stature increase have been

F. J. Rühti - Osteometric Variatíon of the Human Spine 98 two indices for the trarisition period from Late Würm to Post Würm times in males' Most of the robusticity indices for males increased between the Upper Paleolithic and

Mesolithic period, which is not the case for females. For females, between Pre-Würm and

Late Würm time there was ànincrease in humeral robusticity, up by more than 6Yo, and a slight decrease in femoral robusticity, whereas from Late Würm to Post-Würm both robusticities decreased (Jacobs, 1985b). Jacobs (19S5b) describes a decrease in individual male body size mostly within the Upper Paleolithic period and not at the transition to the

Mesolithic time, whereas females showed a continuous reduction. Also body proportions changed during the transition period from Pre- to Post- Würm times in Europe with humerus relative to stature becoming smaller for either sex (Jacobs, 1985b). Limb proportions in Europe did not change according to Frayer (1981). General limb reduction was more prominent for males, explainable by the higher impact of altered hunting conditions, with an 8.8olo decrease for male humerus,7.5o/o for female humerus, 7 -60/o fot male femur and 4.5Yo for female femur, respectively. General stature reduced towards

Mesolithic with 5.5% for males and 3.4Yo for females. Only with the start of the

Mesolithic, at least for males, the stature changed signif,rcantly, while being mostþ stable

for the major Paleolithic periods (Frayer, 1981). Jacobs (1985b) found no such expected

decrease of upper limb robustícily, due to the introduction of the atl-atl and bow and

arïow, between Late Upper Paleolithic and Mesolithic in males, but describes one in

females. Thus, Jacobs (1985b) explains these skeletal alterations to be more linked to

nutritional changes and climatic adaptations, possibly towards a colder environment, than

resulting from technological changes only'

Sexual dimorphism is an important measure to evaluate the ongoing interactions

between a particular environment and the body morphology. Frayer (1980) addressed in

F. J. Rühl¡ - Osteometric Variation of the Human Spine 100

For the particular situation of Europe, since the Neolithic, it has been found that mostly decreasing levels of natural selection and further similarities of cultural environments, rather than migration and its linked gene exchange only, lead to an increased intra-group and decreased inter-group variability in morphological traits

(Henneberg et al., I 978).

Microevolutionary changes, occurring in short well-defined historic time periods, have been shown for various anatomical characteristics e.g., the increase of incidence of the median artery of the forearm (Henneberg and George, 1995), the occurrence of hlperostosis frontalis interna (Hershkovitz et a\.,1999; Rühli and Henneberg, 2002) ot presence of non-osseous tarsal coalitions (Rühli et a\.,2003). Microevolutionary trends as expressed in their significant morphological changes, within short periods of time even question the understanding of modern human origin such as the replacement hypothesis, or the validþ of any taxonomic definition of modern humans in terms of objectively measurable characteristics (Henneberg, 200 1 a).

Surprisingly, microevolutionary changes of the spine seem to be a neglected

research area (Jankauskas, 1994). Some possible secular trends in frequency of spinal

pathologies have been reported, such as the increasing prevalence of spina bifida occulta

(Henneberg and Henneberg, 1999) or the prevalence of spondylarthropathy in baboons

(Rothschild and Rothschild, 1996). Larsen (1980; 1981; 1982) reported a significant

decrease in degenerative spinal joint diseases linked to a cultural shift towards

agricultural lifestyle in an American coastal region, whereas Minne et al. (1988) discuss

in their X-ray based study the influence of the secular increase in body height in the last

century and its impact on spinal morphometry. They describe an increase of vertebral

102 F. J. Rühli - Osteometríc Variation of the Human Spine body height, for the last 110 years and for Th4 to L5 only, of 86 mm, with no alterations,

at least relative to their standard vertebra atTh{.

Nevertheless, according to Jankauskas (1994), there is a lack of microevolutionary

and inter-populational studìes'of the human vertebral column. Just very limited spinal

microevolutionary approaches have been published so far. Secular trends of vertebral

body size (Clark et ø1.,1985; Jankauskas, 1994) or neural canal dimensions (piontek and

Budzynska, 1972; Clark et al., 1985; Tatarek, 2001) have been so far investigated on

lirnited samples only.

Furthermore, according to Jankauskas (1994), no clear definition of the human

spinal osteometry and its variability exists. This is in particular striking since

microevolutionary trends for other major body parts such as skull size (Henneberg, lggg;

Henneberg and Steyn, 1993; Ross and Henneberg, 1995) or stature and postcranial

skeletal dimensions (Schwidetzþ, 1962; Frayer, 1980; Larsen, l9g0; Larsen, lggl;

Larsen, 1982; Formicola, 1983; Frayer, 1984; Jacobs, 1985a; Ruff, 1994; Formicola and

Giannecchini, 1999) have already been addressed in a plethora of reports.

As other possible causes of recent secular trends, genetic factors or their products

acting during early stages of ontogeny, most likely in utero, have been suggested

(Henneberg, 2001b). Furthermore, Henneberg (200lb) names vaccines, or food

containing chemical products interfering with individual growth as additional possible

underlying origins of this secular trend in the most modern times.

To summarize, surprisingly no secular trend of the non-pathologic vertebral

column has so far been widely studied. Hitherto, in the most similar studies, Tatarek

(2001) just focused on the lumbar levels, while Jankauskas (lgg4) included not only a

limited particular Eastern European area, but also choose temporally limited samples

F. J. Rühli Osteometríc - Variation of the Human Spine 103 from the 1" and 2"0 millennium A.D. only. Both studies (Jankauskas,1994; Tatarek, 2001) were, additionally, small in number of spinal measurements taken on each individual.

No investigation focusing on microevolutionary issues on all major levels of the human vertebral column and consisting of a sample dating back to European Late

Pleistocene has been published so far. Furthermore, a combined anthropological and clinical perspective including the morphometric spinal variation as well as the influence of sex and individual age on it, in particular in such a historic sample, has never been fully explored before.

F. J. Rühli - Osteometric Variation of the Human Spine 104 Aim of the study

The aim of this study is to assess and interpret osteometric measures of a number of human spinal landmarks on all major vertebral levels that is cervical, thoracic and lumbar, in Central-Western European skeletal samples dating from the Late Pleistocene to most modern times. The data will be explored with a particular focus on the influence of sex and individual age as well as possible underlying secular and microevolutionary trends. Possible clinical implications will be addressed too.

Hypothesis to be tested

The purpose of this study is to test the null hypothesis that there is no significant change in selected osteometric traits of the human spine in terms of sex and individual age as well as from the Late Pleistocene to modern times in Central-Western Europe.

F. J. Rühli - Osteometric Variation of the Human Spine 105 Material

Dry vertebrae of 348 individuals of both sexes have been included into the study; see also Table 3 for the list of selected individuals or samples, for the complete set of original data see appendix 2 and for apublished abstract onthe data of the present study

see appendix 15. Selection criteria for samples were primarily being of Central-Western

European origin and providing easy accessibility. A list of major samples represented

could be found in Figure 3. The accessibility \Ã/as usually achieved through personal

consent from the collection curator, who also mostly supplied main references and the

collection list, with recorded individual sex and estimated age of the chosen skeletons'

Only unarticulated vertebral columns were used. In case of fragmented bones, only those

whose reconstruction could be done without any apparent size or shape alterations have

been selected.

A1l major historic time periods in Europe since Late Pleistocene are represented,

with the exception of Iron Age and Roman period, when body cremation was the most

popular burial practice in Europe (Schwidetzþ, 1972; Schwidetzky and Rösing, I976).

Years before present (BP) were calculated from 2000 A.D. backwards. The whole sample

(Figure 4) was divided for selected data analysis in three major time groups (Figure 5),

Neolithic I Bro¡g;e Age, Medieval and Modern, respectively. By doing so, the single

individuals from Paleolithic and Mesolithic times were neglected.

The major time periods for Central Europe background are assumed as follows,

mostly according to Straus (1995):

Pleistocene Middle Paleolithic 100,000 - 40,000 B.C'

Late Paleolithiç 40,000 - 10,000 B'C.

Early Upper Paleolithic until 30,000 B'C'

F. J, Rühli - Osteometric Variation of rhe Human Spine 106 Middle Upper P aleolithic 30,000 B.c. - 20,000 B.c.

Late Upper Paleolithic 20,000 B,c. - 10,000 B.c.

Holocene Mesolithic 10,000 B.c. - 4500 B.c.

2000 B.c. Neolithic . ¡ 4500 -

Bronze Age 2000 - 800 B.c.

Old Iron Age (Hallstatt) 800 - 500 B.c.

New Iron Age (La Tène) 5008.c.-04.D.

Roman 0 A.D. - 400 A.D.

Early Medieval 400 - 900 A.D.

Classic Medieval 900 - 1100 A.D.

Late Medieval 1100 - 1500 A.D.

Modem Times after 1500 A.D.

Individual age was known for each skeleton of the "St. Johann" and "Geneva"

provided samples. For the other samples, individual age was recorded based on the

to collection lists. For most of the data analysis individuals were categonzed, according

(20-39 years of age), their estimated core age range, into the three main age groups: adult

(Figure the mature (40-59 years of age) and senile (60 years and older), respectively 6). If

group, core age range of an individual covered more than one major age the individual

group. For was fractioned into these groups according its likelihood to be within each age

counted as example, an individual with the assumed core age of 20-50 years would be

0.67 in the adult and 0.33 in the mature age group'

The geographic background of the selected samples was from Southern Germany,

Switzerland, Austria and France. A geographic overview of the origin of the samples

could be found in Figure 7.

107 F. J. Rühti - Osteometric Variation of the Human Spine Table 3 Individuals / samples included in the present study

SAMPLE / N-SEX YEARSBP CURRENTLOCATION SELECTED SPECIMEN (TOTAL: ì REFERENCES L79m,l69f)

La Ferrassie I 1-m 30,000 Musée de I'Homme, Paris (OaKey et al., (Homo søpìens (France) I97l;Heim,1976; neøndertalensís) Sfrnger et al., le84)

La Chapelle-aux- I m 30,000 Musée de I'Homme, Paris (Oahey et al., Saints I 1971; Stringer e/ (Homo søpiens al.,1984; Trinkaus, neøndertalensis) le85)

Cro-Magnon 11 2 1 - m, I - f 25,000 Musée de I'Homme, Paris (OaKey et al., l97l; Stringer e/ al.,1984)

AbriPataud6 l-m 18,250 Musée de I'Homme, Paris (OakTey et al., l97l; Stringer e/ al.,1984)

Neuessing 1-m 18,200 Anthropologische (OaYJey et al., Staatssammlung, München l97l; Schröter, (Germany) r977)

Veyrier 1-m 12,000 Département d'Anthropologie, (Pittard and Sauter, Université de Genève 1945; OaKey et al., (Switzerland) r97r)

Le Bichon l-m I 1,700 Latènium, Hauterive (Sauter, 1956; (Switzerland) Oakley et al.,l97l; Morel, 1993)

Gramat I 1-m 8000 Institut de la Paléontologie (Newell e/ a/., Humaine, Paris 1979;Bodenet al., le90)

Yaihingen/Enz 4-m,5-f 7200 Anthropologisches Forschungsinstitut, Aesch (Switzerland)

Wandersleben 15 -m,26-1 7000 Zentrum Anatomie, Georg- (Carli-Thiele, 1996) August-Universität, Göttingen (Germany)

F. J. Rühli - Osteometric Variation of the Human Spine 108 Iloëdic 8, 9 1-m,1-f 6600 Institut de la Paléontologie (Vallois and de Humaine, Paris Félíce, 1977; Newell et a\.,1979)

Téviec1,16 1-m,1-f 6600 Institut de la Paléontologre (Newell e/ a/., Humaine, Paris 1979;Bodenet al., 19e0)

Birsmatten l-f 6300 KantonsmuseumBasel-Land, (Sedlmeierand Liestal (Switzerland) Kaufmann, 1996)

Hainburg l7 -m,23-f 3800 - 3500 Naturhistorisches Museum, V/ien (Ehgartner' 1959) (Austria)

Straubing 37 -m,44-f 1500 - 1300 Zentrum Anatomie, Georg- (Kreutz, 1997) August-Universität, Göttingen

Aesch 9-m,5-f 1370 - 1300 Anthropologisches Forschungsinstitut, Aesch

Barbing 15-m,12-f 1300 Zentrum Anatomie, Georg- August-Universität, Göttingen

Winterthur 23-m,13-f 950-435 AnthropologischesInstitut, (Jäggietal.,1993) Universifät Zürich (S witzerland)

Chur 8-m,7-f 750-550 A¡thropologisches Forschungsinstitut, Aesch

St. Johann 20-m,17-f 228-163 Naturhistorisches Museum, Basel (Etter, 1988; Etter (Switzerland) and Lörcher, 1993)

ttGênevatt* 5-m,4-f 135-80 Département d'AnthroPologie, Université de Genève

otGenevatt* 9-m,4-1 133-80 Département d'Anthropologie, Université de Genève

ttGenevatt* 5-m,3-f 120-66 Département d'Anthropologie, Université de Genève

ttGeneYat'* 2-m,2-f 106-85 Département d'AnthroPologie, Université de Genève

* víllages Apples, Bex, La Sarraz and Saint-Prex summarizedfor reasotts of anonymíty

109 F. J. Rühli - Osteometric Varíatíon of the Human Spíne Major samPles (N t 5)

50 44 E8 males 45 trfemales 40 37

35

30 26 N25 23 23 20 21 20 17 1 15 13 13 15 12 9 10 87 45 5 5

0 ct o) o) 5 c (d 0) c o c. o) f Ø (ú c) -o E o) õ Ë o -c '= (ü o o c) 'õ 'õ (ú Íl c ? (, I U) U) =

Figure 3: Major samPles examined

Time groups E males 1 00 92 trfemales 90 81 80 70 60

N 50 41 40 34 30 30 22 23 17 20 7 10 1 0 Paleolithic Mesolithic / Bronze Age Middle Ages Modern Neolithic

Figure 4: Historic age groups (time groups) represented in the whole sample

110 F. J. Rühli - Osteometric Variation of the Human Spine Major time grouPs K males Elfemales 100 92

90 81 80 70

60 54

N50 41 36 40 30 30

20

10

0 Neolithic / Bronze Age Middle Ages Modern

Figure 5: Major time groups represented in the whole sample

Age grouPs

120

96.83 Emales 100 86.57 trfemales

80 66.1

N60 48.65

40 26.33 23.53

20

0 Adult Mature Senile

Figure 6: Age groups distribution in the whole sample

111 F. J, Rühti - Osteomelric Variation of the Human Spine Supplementary information on the samples

The La Ferrassie I individual is considered an adult male Neandertal of approximately 40-50 years of age. He was discovered in 1909 in Savignac du Bugue, 40 km southeast of Périgueux in the Dordogne Region of France and most likely dates to the Würm II period. A reference list of the suggested chronostratigraphic dates of the various Würm periods can be found elsewhere (Smith, 1984). His spine is of general high robusticity similar to the one of the La Chapelle-aux-Saints 1 Neandertal individual. It represents one of the most complete preserved Neandertal vertebral columns (Oakley et al., 1971; Heim,1976; Stringer et al., 1984; Riel-Salvatore and

Clark, 2001).

The La Chapelle-aux-Saints 1 individual, a holotype of the Homo chapellensis and supposed to be a Neandertal, was found in 1908 near Corrèze in Central-Southern

France. The male adult individual of approximately 40-50 years of age is linked to the

Würm II period. This individual was supposed to be 164 cm in height and of a body weight of 70 kg. The vertebral column of this individual drew attention in earlier times, but the view of his apparently primitive anthropoid-like neck is less supported nowadays, as it was in the times after its discovery (Stewart, 1962; Oakley et al.,I97l;

Stringer et al., 1984; Trinkaus, 1985; Rufl 1994; Riel-Salvatore and Clark, 2001).

Despite its arthritic changes of the cervical and thoracic spine (Trinkaus, 1985), this individual was included into this series due to its historic importance.

The Cro-Magnon individuals, anatomically modern Homo sapiens, were discovered in 1868 near the station Les Eyzies de Tayac, approximately 25 km northwest from Sarlat in the Dordogne Region in France and date to the Würm III

F. J. Rühli - Osteometric Variation of the Human Spine 112 period. Cro-Magnon 1, also called "the Veillard", is the holotype of Homo spelaeus and

!ù/as supposed to be a male of at least 45 years of age. Cro-Magnon 2 is believed to be an adult female of approximately 20-30 years (Oakley et ø1., I97l; Stringer et al., 1984;

Riel-Salvatore and Clark, 2001).

The Abri Pataud 6 individual, apparently an adult male, was excavated in 1963

in Les Eyzies, 25 krrL north-west of Sarlat in the Dordogne Region in South-Western

France (Oakley et al.,l97l; Stringer et al.,1984).

The Neuessing 2 individual was found in l9l3 in the Altmühl Valley,

approximately 25 km southwest from Regensburg, Southern Germany. It was dated to

the Weichselian-Wirm period and is supposed to be an adult male individual, of

approximately 30 years (Oakley et al.,l97l)'

The Late Paleolithic Magdalenian type, Late Würm period, Veyrier skeleton was

discovered in 1916 in Veyrier in the Haute-Savoy region in France, next to the actual

Swiss border. His living stature is estimated to be 169 cm, which makes him shorter

than the average Cro-Magnon humans, but still larger than the Magdalenians and most

'Westem European Mesolithic and European Neolithic people. Humeral and femoral

robusticity are both small (Pittard and sauter,1945; Oakley et al.,l97l).

The Le Bichon individual, which was found in a cave at an altitude of

approximately 850 meters above sea level in 1956 next to La Chaux-de-Fonds, Western

Switzerland, is the oldest preserved individual of nowadays Swiss geographic

background and belongs to the Late Paleolithic Cro-Magnon type. His cause of death, as

a side remark, was recently reconstructed to be a hunting accident (Morel, 1993).

F. J. Rühli - Osteometríc Varíation of the Human Spine 113 Although that his stature could not have been completely reconstructed, the individual was apparently not very tall (Sauter, 1956; Oakley et al.,l97l)'

Hoëdic is a Mesolithic site of nine adult individual skeletons, excavated in the

1930's and located on a small island at the Bretagne, on the North West coast of France.

Téviec is a Mesolithic series, possibly slightly older than the Hoëdic site, consisting of 15 adult individuals situated in similar environment, and32 km further in

North East direction. This site was excavated primarily in the late 1920's. Both islands,

Téviec and Hoëdic, were supposed to be even easier accessible today than in the

Mesolithic, most likely by a dry walk from the mainland. The Hoëdic individuals seem to be not of massive robusticity, which is similar to the Téviec individuals. Apparently in one of the nine Hoëdic graves, an individual was found with six lumbar vertebrae. In addition, the Téviec sample contains one individual with such an increased number of vertebrae. The individuals from both Mesolithic samples are of small stature, at least in comparison with East Europeans of the Late Paleolithic, but they are comparable to other Mesolithic people of similar geographic background. Individual stature was for the

Hoëdic males on average 160 cm and for the females I52 cm, whereas it was 159 cm for the Téviec males and 151 cm for females, respectively. The two samples were classified to be modern humans of the "Téviec-island" type (Vallois and de Félice, 1977).

In comparison, the Gramat male from mid-South-Central France was reconstructed to be of 165 cm height and of a remarkable femoral robusticity, but not of high humeral robusticity: He seems to be an exceptional human of the "Téviec- continental" t¡pe (Vallois and de Félice, L977). The Gramat male individual is a complete skeleton of the Holocene period discovered in 1928 in Le Cuzouln de Gramat,

F. J. Rühli - Osteometric Variation of the Human Spine 114 approximately 55 km north-east of Cahors in the Dordogne region of France (Oakley e/ al., I97l). For a precise description of the distinctive skeletal characteristics of the two

Mesolithic prototypes, "Téviec-continental" and "Téviec-island", see Vallois and de

Félice (t977).

The Holocene Birsmatten individual, most likely to be a female according to ne,w anthropological assessments, was found in 1944 in Nenzlingen (Northern

Switzerland) and is the only Mesolithic body burial in nowadays Switzerland. The skeleton is of remarkable preservation for its historic age and individual stature was calculated to be of approximately 160 cm (Sedlmeier and Kaufmann, 1996).

Wandersleben is a Neolithic Linienbandkeramiker (linear pottery) - culture

settlement, located between Gotha and Erfurt in present-day Germany. The whole

sample consists of approxim ately 200 individuals, representing one of the largest known

Central European classic settled agricultural lifestyle societies, but an archaeological

report of this excavation is stili not yet published (Carli-Thiele, 1996).

This situation is similar for the sample of Vaihingen, which is also a linear

pottery settlement (Early Flomborn and Middle linear pottery phase) in nowadays

Vaihingen an der Enz, ín the Neckar Region next to Stuttgart (Baden-Württemberg,

Southern Germany) of generally excellent preservation. A final report on this old

Neolithic agricultural site with approximately 100 flexed burials has not yet been

published; preliminary information could be found at the following internet-website:

htþ ://home.bawue. del-wmwerner/gtabung/vaih.html.

Hainburg is a burial ground of 253 skeletons from the Early Bronze Age

Wieselburger - culture, excavated in the late 1920's as well as in the late 1930's. The

F. J. Rühli - Osteometric Variatíon of the Human Spine 115

four Swiss villages with a mostly listed age at death, sex and profession. They lived in

background e'g'' craftsmen or farming background, but some had a similar professional

sample' light industrial workers, as the individuals of St' Johann

117 F. J, Rühli - Osteometric Variation of the Human Spine

Methods

Measurements

Osteometric measurements were taken on the following vertebral levels

(numbered as counted from cranial):

- 3'u (CERVICAL 3 vertebra in a normal spine with 24 pre-sacral vertebrae)

- 7'n (CERVICAL 7)

- B'n (THORACIC 1)

- 13'n (THORACTC 6)

- t7" (THoRACrc 10)

- 20* (LUMBAR 1)

- 24" (LUMBAR 5)

The vertebral levels were selected for the following reasons:

C3 is the f,rrst cranial vertebra with a true vertebral body; therefore, it acts as a

transition vertebra between the cranial base / upper cervical spine and the main

cervical spine

C7 also called vertebra prominens due to its outstanding spinous process; it is the

last vertebra of the cervical spine, therefore, acts as a transition vertebra between

two of the major spine sections

Thl is the fìrst thoracic vertebra; transition vertebra between the cervical and thoracic

spine

F. J. Rühli - Osteomelric Variation of the Human Spine 719

A1l measurements were done on original specimens only. One single observer took all vertebral measurements, so no inter-observer error occulTed. All measurements were taken twice repeatedly. If the results showed a difference of more than 0.1 mm a third measurement was performed and the average of all assessments was later used for analysis. Any bones manifesting major gross morphological abnormalities e,g., severe arthritic changes on multiple levels or diffuse idiopathic skeletal hyperostosis, were excluded. If any bone was fractured, only the ones allowing perfect re-adaptations of the broken pieces were assessed. If one side of the transverse process was missing but the

other side was preserved intact, overall transverse process width was estimated by

multiplying by the factor of two the distance from the intact most lateral tip to the

middle of the endplate at the posterior border of the vertebral body. Minor osteophytic

alterations were not a reason for exclusion, as long as they were regarded as normal age-

related adaptations. Young adult individuals showing macroscopic signs of still ongoing

vertebral growth were excluded.

To assess the suggested osteometric variation and possible microevolutionary

trends of the human spine, a set of various measurements was performed at each chosen

vertebral level; see also Table 4 and for all abbreviations used see appendix 1. In

accordance with most of the earlier published major studies dealing with spinal

morphometry, such as e.g., the ones by Jankauskas (1994) or Panjabi et al. (l99la;

1991b; 1992), dimensions of various anatomical parts of the selected vertebral levels

were chosen. To determine potential alterations of the vertebral bodies, measurements

of their height and main diameters were performed. To be able to detect likely

alterations of the pedicles, the maximum pedicle height, was included as well. For the

assessment of the osseous outline of neural pathways, the main diameters of the spinal

F. J. Rühti - Osteometric Variation of the Human Spine 121 canal, the foramen magnum as weil as the width of the intervertebral foramen were chosen to be measured.

Length and circumference of the femur and humerus were included for the assessment of individual stature as e.g., shown by Trotter and Gleser (1952) and robusticity, as already outlined e.8., by Martin (1928). One has to be aware that humerus maximum length and radius maximum length as well as femur bi-condylar length and

tibia maximum length are strongly correlated not only in recent samples, but also in

Neandertals and early anatomically modern humans (Trinkaus, 1981). Therefore, all

findings in the measured long bones may also be generally true for the other related

limb bones. Furthermore, Martin (1928) already dehned the measurement of femoral

head width and bi-iliac width, both indicators of individual body mass as e.g., applied

by Ruff er al. (1997) for Pleistoc ene Homo and used in the study presented here as well.

Most of the selected measurements were performed according to the well-

established osteometric definitions by Martin (1928). Martin (1928) did not define

osteometric measurements for e.g., maximum transverse process width or spinous

process length. The first one was done in the present study according to Hasebe (1913)

and the latter one according to Schultz (1961). The maximum pedicle height was

dehned hereby similar as in the study by Shapiro (i993). Furthermore, a plethora of

definitions for the measurement of the intervertebral foramen width and height,

especially for cadaveric samples, has been dehned so far; see also Figures 8 and 9 with

unaltered or slightly adapted figures of earlier publications. In the present study, a

measurement approach similar to the ones chosen by Amonoo-Kuofi (1985) or

Ebraheim et al. (1997), was performed.

F. J. Rühli - Osteometric Variatíon of the Human Spine 122 Table 4: Measurements used (M: numbering according to Martin 1928)

t Abbreviation Ventral cranio-caudal diametei of vertebral body: Ml Dorsal cranio-caudal diameter of vertebral body: M2 Mean sagittal diameter of vertebral body: M6 Mean transverse diameter of vertebral body: M9 Maximum pedicle height; see also Figure 8: PH (Shapiro,1993) (Hasebe, Spinous process length: SPL 1913)

Transverse process width: TPW (Schultz,l96l) cranial / caudal intervertebral foramen width; see also Figure 8 / 9: IFCR / IFCA (Amonoo-Kuofi, 1985;

Ebraheim et al., 1997)

Sagittal diameter of vertebral foramen: M10

Transverse diameter of vertebral foramen: Ml1

Foramen magnum breadth: FMM16

Foramen magnum length (bøsion - opisthion): FMMT Maximum length of humerus: HLMI Minimal circumference of humerus: HCMT Maximum length of femur: FLM1 Circumference at mid-femur: FCM8 Femoral head breadth' FHM18 Bi-iliac width: BIWM2

123 F. J. Rühli - Osteometric Variation of the Human Spine

DM

UËþ.. \trrh¡õ I hldrtrbñi dÈ I ( .t l,í tl

L0# trttafr¡

H G

J I

Fig. 8 (cont.): Lateral views of measurement definitions of the intervertebral foramen

G) panjabi et al. (1983); A) area of the notch, h) maximum vertical space, w) minimum width of the foramen H) Ebraheim et al. (1996);DM) medial zone depth I) Cinotti et at. (2002); 1) superior foraminal width, 2) minimum foraminal width, 3) pedicle length ¡; Prãsent study; 1) cranial intervertebral foramen width, 2) caudal intervertebral foramen width 3a) dorsal vertebral body height, 3b) maximum pedicle height; intervertebral foramen dehnitions similar e.g., to Amonoo-Kuofi (1985) or Ebraheim et al. (1997)

F. J. Rühli - Osteometric Variation of the Human Spine 725 MBdìäl ¿dns

Middle êqñe

Lsiùäl Z6n€

A B

c

Figure 9: Cranial views of osteometric measurement dehnitions of the intervertebral foramen

A) Inufusa et at. (1996): 1) mid-sagittal diameter of vertebral canal, 2) sub-articular ligamentum flavum thickness sagittal canal diameter, 3) 'WL) B) Ebraheim et al. (1996): WM) medial zone width, lateral zone width, D) distance from vertebral body midline to the anterior border of the medial zone, A) angle between the nerve groove axis and the mid-sagittal plane C) Present study: 1) intervertebral foramen width (similar for cranial and caudal measurement)

F. J. Rühli - Osteometric Variation of lhe Human Spine 126 Symmetrical structures, such as the intervertebral foramen, were measured bilaterally; since, according to Marchesi et al. (1988), at least some of the vertebral measurements show side-dependent values of unknown signihcance.

Long bones measures were taken preferably on the right side, if preservation allowed it. Martin (1928) stated that the right humerus is usually longer and more massive than its left counterpart is. Therefore, the right sided long bones were chosen in this study, despite the fact that some authors use the left side to assess postcranial dimensions (Larsen, 19Sl). The left femur is usually bigger than the right one, whereas it is the other way round for the humerus (Martin, 1928; Trotter and Gleser, 1952).

According to pfeiffer (1980), the long bones of the non-dominant side, which is usually the left one, are less susceptible to age dependent size and robusticity changes.

Nevertheless, correlations between right and left side measurements of long bones are

high. According to Trotter and Gleser (1952), in white males inter-correlation among

lengths of right and ieft femur as well as humerus is for both long bones 0.98, with

mean absolute side differences for femur and humerus 0.6 mm and 0.5 mm respectively'

Either long bone measurements were performed in the study presented here by the

author himself, to the nearest 1 mm, or they were taken from collection references.

paleolithic long bone and foramen magnum data were kindly provided by

other earlier published data (Martin' Holliday (T. Holliday , pers. comm.) or gained f¡om

l92g; Trinkaus et at.,1994). Sex and age of Paleolithic and Mesolithic skeletons were,

in addition to the listed main references, brought in accordance with various sources

(Holliday, 1997; Formicola and Gia¡necchini,1999; Holliday, 1999; Riel-Salvatore and

127 F. J. Rühli - Osteometric Variation of the Human Spine Clark, 2001). Individuals of the Hainburg sample were assed in terms of sex, based on new estimations (B. Auerbach,p ers. comm.) or collection references (Ehgartner, 1959).

Furthermore, the measurements used for the sagittal and transverse vertebral body diameter taken at mid-height escape most of the degenerative lesions, since these pathologies appear preferably at the level of the superior or inferior endplates. On Th6 and Th10, no cranial intervertebral foramen width could be determined. With regard to the particular anatomy of the posterior surface of the vertebral body (Larsen, 1985), it is worth noting, that in this study the sagittal vertebral body diameter was measured according to Martin (1928). In the midline of the posterior surface, the bridge of the foraminae caused by the basivertebral veins was the posterior reference point for the diameter. This point does usually slightly differ from the most concave point within the posterior surface at least of the lumbar vertebrae (Larsen, 1985).

Technical equipment

All measurements, except for long bone length, circumference and bi-iliac

breadth, were taken with a sliding caliper to the nearest 0.1 mm.

Several authors (Ktag et al., 1988; Scoles et ø1., 1988) pointed out that direct

To osteometric measurements are still the best method to determine spinal dimensions'

improve direct caliper based measurements, Ebraheim et al' (1996) e'g', even cut off the

transverse process of the particular level. This would not be favoured for obvious

reasons in historic specimens. Surprisingly, Yoo et at. (1992) state that a caliper-based

due to assessment of the intervertebral foramen diameter is not accurate enough, mainly

the measurement technique itself. Therefore, they used for their study of intervertebral

128 F. J. Rühli - Osteometric Variation of the Human Spíne foramen size in fresh but frozen cervical cadaver spines a penetrating probe. Obviously, this would not useful for skeletal studies either.

Es timation oJ' intra- ond ini"r- observer error

It is crucial to know a possible intra-observer error of osteometric measurements of the vertebral column. In average this my have an extent of approximately 0.25 mm per vertebra (Todd and Pyle, 1928a; Lanier, 1939). Larsen (1985) addressed the possible e¡¡or caused by an uneven vertebral surface with a possible error of up to several tenth of a millimetre. Nissan and Gilad (1986) found in their caliper based roentgenogram study, that for osseous vertebral measurements, statistical errors are of higher importance than the measuring linked effors. Nissan and Gilad (1984; 1986) observed the intra-observer error in defining skeletal landmarks in a radiological study to be of

0.5 mm or less. The average intra-observer etïor of measurement for a semi-automatic

measurement of vertebral dimensions in conventional radiography was l.4Yo, arrd the

inter-observer erïor was 2.lo/o (Diacinti et al., 1995). Kandziora et al. (2001) describe

the error of osteometric measurements of the cervical spine to be +/- 0'08 mm in their

study by using a digital ruler with a stated accuracy of 0.1 mm. They found an equal

accuracy of the radiologic assessments. Hinck et al. (1966) describe the intra-observer

error in an X-ray study of the interpediculate distance to be of 0.26 mm. Furthermore,

Minne et al. (1988) report a low intra- and inter-observer effor of measurement in their

X-ray study on the normal spinal morphometry. Todd and Pyle (1928b) discussed the

extent of erors between roentgenographic and wet spine morphology, as well as the

intra-observer error of measurement on dry and wet spines (Todd and Pyle, 1928a).

Roaf (1960) found an acceptable correlation between radiographic and post mortem

F. J. Rühli - Osteometric Variatíon of the Human Spine 129 spinal measurements. Jacobs (1985b) lists an intra-observer error of 0.002Yo for lengths and l.lYo for other measurements. In comparison to earlier published data, his margin of effor was 0.003% añ,Z.IYo respectively. Therefore, Jacobs (1985b) concludes that by including published data in âpersonaliy acquired data sample, one does not signihcantly increase existing intra-observer errors of measurements. On the other hand, Porter et al.

(1937) had in their ultrasound based study a mean repeatability in measuring the 15" oblique lumbar spinal canal width of 0.5 mm. ln another ultrasound based study of the oblique lumbar spinal canal dimension, Porter et al. (1978b) found an intra- and inter- observer error of measurement of 0.2 mm. The intra-observer and inter-observer error of measurement were both 0.4 mm in another ultrasound study of the same structure

(Hibbert et al., 1981a). For a similar study, Legg and Gibbs (1984) report an intra-

observer error of measurement of less than 0.3 mm. Surprisingly, they had consistently

different values obtained than earlier published ultrasonographic assessments of the

spinal canal diameter (Porter et a1.,1978a), explained by them to be most likely due to a

systematic difference. The intra- and inter-observer coeffrcients of variation were

approximately 25Yo atd 5o/o, respectively, in an X-ray based morphometric study by

Hermann et al. (1993). Furthermore, they mention the possible error in different X-ray

studies caused by the fact that average subcutaneous fat thickness in selected

populations varies and, therefore, by having an altered magnification factor while

obtaining the X-rays, the gained data may differ slightly as well. Additional technical

factors relevant especially for radiographic studies of spinal morphology are also

mentioned by Hermann et al. (1993). In their anatomic-biomechanical study on the

cadaveric lumbar spine, Fujiwara et øt. (200I) determined the intra-observer error for

130 F. J. Rühti - Osteometric Variation of the Human Spine the measurement of the intervertebral foramen width to be 0.3 mm - 0.4 mm and for the intervertebral foramen height to be 0.2 mm. The inter-observer error is well known for spinal measurements in clinical imaging situations (Ullrich et al., 1980; Beers et al.,

1985; Galla gher et at., tggg;Hermann et a1.,1993; Wildermuth et a1.,1998), but this does not apply for this study due to the fact that only one observer performed all measurements. Ullrich et al. (1980) list the inter-observer effor for linear spinal measurements by CT to be of less than 3Yo. Gallagher et a/. (1988) examined the intra- and inter-observer error of measurement in a radiographic study on female spines. They found variation coefficients to be of less than 3Yo or 4To, respectively, for linear vertebral measurements.

The standard error of measurement for the pedicle length, as measured by

Zindrick et al. (1987) in a radiographic measurement was for the thoracic and lumbar spine between 0.2 and 0.6 mm. For a slightly different way of osteometric measurements of the pedicle height, Marchesi et al. (1988) found standard errors of measurements between 0.2 mm and 0.4 mm; for the osteometric assessment of the spinal canal dimensions errors of 0.2 mm - 0.7 mm, and for the anterior and posterior vertebral body height errors between 0.2 mm and 0.5 mm. Olsewski ¿/ al. (1990) mention an error of measurement for pedicle height and width of 0.1 mm. Kothe et al.

(1996) found an accuracy for the digitised measurement of pedicle slices to be 0.06 mm.

Misenhimer et al. (1989) describe the accuracy of CT measurements of the pedicle in comparison to osteometric data to be within a third of a millimetre. Panjabi et al.

(L99la; 1992) list in their three-dimensional morphological studies, which are largely different from the one presented here, the overall error in computing vertebral

F. J. Rühli - Osteometric Variation of the Human Spine 131

2.3 otr 2.1 oE o5 1.9 (E o 1.7 E o 1.5 o 1.3 o (! 1.1 .9 c 0.9 .C(t o 0.7 0.5 Experienced lnexperienced 1 lnexperienced 2 lnexperienced 3 Observer

Figure 10 Technical error of measurement for selected spinal dimensions by experienced and inexperienced observers

F. J. Rühli - Osteometric Variation of the Hunan Spíne 133

Examination permis si on

Collection accesses were approved in oral or written form by the responsible

curators in advance.

Data analysis

All original data were copied by the author himself into a Microsoft@ Excel 'WA, 2000 (Microsoft Corporation, Redmond, USA) spreadsheet. Data were checked

twice for obvious mal-transcription errors. If any doubt about data persisted after

double-checking with the original record sheet, the particular measurement was deleted,

It can be assumed, based on earlier reports (Minne et al., 1988; Black et al.,

1991; Xu et al., 1995), that spinal morphometric ratios follow a normal or Gaussian

distribution. Therefore, measurements before the final data analysis were trimmed by

deleting all data outside the range ofthree standard deviations. Spinous process length

on C3 (C3S1) and transverse process width on level C7 (C7Hl) were excluded in most

data analyses due to their overall small sample size.

Statistical analyses were done by either using Microsoft@ Excel 2000 or,

primarily SPSS@ 11.0 (SPSS Inc. Chicago, IL, USA) software. The skeletal sample was

analysed separately for both sexes. The limits of two-tailed significance were estimated

for p<0.05, with Bonferroni's correction added for measurements on multiple vertebral

levels. Morphometric values were listed including means and standard deviations as

well as mode, median and minimum and maximum values. Standard deviations for

Table 6 were calculated as sample standard deviations, whereas for the data sets in the

appendices it was defined as population standard deviations. Sexual dimorphism of

measurements was assessed as a percentage difference of mean values as well as by

F, J. Rühti - Osteometric Variation of the Human Spine 135 paired t-test. Paired t-test was also applied for analysis of side differences of spinal measurements. Correlation of variables with individual age was tested primarily on the well-recorded modern samples. Furthermore, correlations of variables with major age groups, dehned as adult, mature and senile, were tested for the non-modern samples, as well as for the three major time groups, defined as Neolithic I Bronze Age, Medieval and modern, respectively. Temporal trends were considered for the whole sample, including the single individuals from the Mesolithic and Paleolithic time period. To test for the best regression model, linear, quadratic, cubic, exponential, logarithmic and power functions were assessed. One-way analysis of variance (ANOVA) was used to test for signifrcant alterations of mean values and standard deviations of variables between the three major time groups. Principal components analysis was done for the whole sample, separately for both sexes.

A list for all used abbreviations for the spinal variables could be found in

appendix 1.

Critical sample size

The critical sample size to detect morphometric measurements depends on the

level of significant mean difference (E) between samples. It is

E-: SD/{N

with 2'E,:mean critical difference, SD being the standard deviation, and N the number

of individuals.

If a difference of */- one SD is expected, the critical N should be 4. If a

difference of a half of SD is expected a critical sample size of 16 and with a difference

of a third of SD it is 36 and with a quarter SD it is 64. A discrepancy of +/- one SD is a

F. J. Rühti - Osteometric Varialion of the Human Spine 136

Table 6: Composition of modern samples St. Johann and "Geneva", with individually known sex and age (N:71, Meat=49.4 yrs, SD:18.4 yrs)

Age group - ',N males N females (Meáñ:51.9 yrs, (Mean:45.9 yrs, SD=I8.6 yrs) SD:l8.3 yrs)

20-39 yrs 13 15

40-59 yrs l4 8

>60 yrs L4 7

Total 4l 30

F. J. Rühli - Osteometric Tørialion of lhe Human Spine 138

L1M6

30

20

10

() Std. Dev = 2.31 c) Mean=27.6 qf c) N = 132.00 LL 220 24.0 26.0 28.0 30.0 32'0 34.0 23 o 2s.0 27.O 29.0 31.0 33 0 3s 0

L1M6

Figure 11: Sagittal diameter of Ll (LlM6) vertebral body in females showing mostly a norÏnal distribution

LsTPW

20

10

() Std. Dev = 15.04 c) ú Mean = 78.0 o N = 70.00 LL 0 100.0 40.0 50.0 60.0 70.0 80.0 90.0 45.0 55.0 65.0 75.0 85.0 95.0

LsTPW

Figure 12: Transverse process width of L5 (L5TPW) in females showing a non- normal distribution

140 F. J. Ri)hti - Osteometric Variation of the Human Spine L1 M1O

30

20

10

() É Std. Dev 1.53 (l) = = Mean = 17.80 oC' II 0 N = 135.00 '""írø'r""r{r"íuø':ojr.¿ieo"'e*ieoier+eo}Ç:oo1Çeoo

L1M1O

Figure 13: Transverse spinal canal diameters at L1 (LlMl0) in males showing mostly a normal distribution

C3SPL

10

6

4

o c Std. Dev = 3.26 c) q Mean = 16.6 o N = 40.00 LL 0

1 1 .0 13.0 1 5.0 17 .0 19.0 21 .0 23.0 12.0 14.0 16.0 18.0 20.0 22.0

C3SPL

Figure 14: Spinous process lengths at C3 (C3SPL) in males showing a non-norrnal distribution

F. J. Rühli - Osteometric Variation of the Human Spine 1.41 Osteometric data of the whole sample

The basic osteometric data, consisting of mean, standard deviation and number of measurements of a particular variable, are presented for both sexes sepffately in

Table 7. Also listed in Table 7 are the same measures for the subgroup of "modern" individuals. Graphs of the examined spinal variables are shown, for the modern subgroups only, in Figure 1 5. The range curves, beside mean graphs for both sexes, are shown. Since usually females have smaller values than males, the female "mean minus standard deviation" is smaller than the same limits in males; therefore, just the female curve is shown. On the other hand, males will have higher values for the "mean plus standard deviation" - curve; therefore, their curve is shown as upper limit of range. As the only exception, in case of intervertebral foramen width the maximum range is defined by female "mean plus standard deviation" and males "mean minus standard deviation". The osteometric pattern for the modem samples is as follows:

The ventral vertebral body height shows generally a consistent increase from C3 caudally to the last lumbar levels in both sexes. The dorsal vertebral body height increases caudally from C3 to Ll and decreases for the last lumbar level in both in sexes.

For the sagittal vertebral body diameter, there is a consistent increase caudally in both sexes. The transverse vertebral body diameter also displays in general an increase caudally, again consistent in both sexes, but with the single exception of Th6, which shows slightly smaller values than Thl.

Pedicle heights follow a similar pattern on both sides and are bigger in males than in females. The pedicle heights show an increase caudally from C3 to Ll, with a decrease caudally in size for the last lumbar level.

F. J. Rühti - Osteometric Variation of the Human Spine 1'42 The diameters of the osseous spinal canal show a different pattern. In both sexes there is a decrease in sagittal spinal canal size from C3 caudally with a subsequent increase in the upper thoracic spine to level Th6. Level ThlO shows slightly smaller values than Th6 in males, but increased means in females. Another increase caudally in the lumbar spine can be demonstrated and, finally, there is a decrease for the last lumbar level. The transverse diameter shows, again consistent for both sexes, an increase from level C3 to C7, followed by a decrease caudally till Th6, with a steady increase caudally; consistent in both sexes.

The spinous process length shows consistent in both sexes increase caudally from C3 to Thl, with a subsequent decrease until Th6 and another increase caudally.

The last lumbar level finally shows a smaller spinous process than Ll.

The transverse process width shows for both sexes an increase caudally in the cervical region, with a decrease for the thoracic levels and another increase in the lumbar region.

The cranial intervertebral foramen widths are bigger than the caudal ones on the same vertebra. The cranial intervertebral widths show consistent in both sexes similar values for most regions, except for level Ll, which shows by far the biggest means.

The caudal intervertebral foramen widths increase in size from C3 caudally till

Th10, again similar for both sexes. Whereas in females, Ll shows bigger values than

Th10, in males the means of the first lumbar level are equal or even smaller than the ones to be found at Thl0. Both sexes show a decrease in size for the last lumbar level.

The foramen magnum demonstrates bigger values in males than in females, with the sagittal diameter being larger than the transverse one. All long bone dimensions, including bi-itiac width, are bigger in males than in females.

F. J. Rühli - Osteometric Variation of the Human Spine 143

30 Dorsal vertebral bodY height 28 Females (mean) 26 ------Females (mean-SD) 24 - (mean)

22 (mean+SD) --Males -Males ÈEzo 18

16

14

12

10 c3 T1 T6 T10 L1 L5 vertebral level

Ventral vertebral bodY height

30 Females (mean) 28 ------26 Females (mean-SD) - 24 (mean) 22 (mean+SD) - -Males E 20 -Males E 18 16 14 12 10 c3 c7 T1 T6 T10 L1 L5 vertebral level

40 Sagittal diameter vertebral body

35 Females (mean) - " "' Females (mean-SD) - 30 (mean)

(mean+SD) E 25 - -Males E -Males

20

15

10 c3 c7 T1 T6 T10 L1 L5 vertebral level

Figure 15: Variables by vertebral levels with mean for males and females and maximum one standard deviation range (male mean+SD, female mean- SD)

F. J. Rühli - Osteometric Variation of the Human Spine 745 Transverse diameter vertebral body 55 50 Females (mean) ------Females (mean-SD) 45 - (mean) 40 (mean+SD) 35 -Males- -Males E E 30 -: - -./ 25 20 15 10 UJ T1 T6 T10 L1 L5 vertebral level

Left pedicle height

18 Females (mean) 16 ------Females (mean-SD) - 14 (mean) (mean+SD) È-12 -Males- -Males Ero

8

6 4 c3 c7 T1 T6 T10 L1 L5 vertebral level

Right pedicle height

18 (mean) -\. \. - - -. -. (mean-SD) -' 16 Females - -Females (mean) 14 (mean+sD) 812 -Males- -Males E 10

8 6 4 c3 c7 T1 T6 T10 L1 L5 vertebral level

Figure 15 (cont.): Variables by vertebral levels with mean for males and females and maximum one standard deviation range (male mean*SD, female mean-sD)

F. J. Rilhli - Osteometric Variation of the Human Spine 1.46 Sag¡ttal diameter sPinal canal 20

19

18 ,4êé4¿ 17 -\r\--¿?-n E re tr 15 ---- Females (mean) ------Females (mean-SD) 14 (mean) 13 r rMales (mean+SD) 't2 -Males c3 c7 T1 T6 T10 L1 L5 vertebral level

Transverse diameter sP¡nal canal 30 28 tt¿ ^ \ 26 24 Ezz E 20 Females (mean) - - -. - - Females (mean-SD) 18 - (mean) 16 (mean+SD) - -Males 14 -Mdes c3 c7 T1 T6 T10 L1 L5 vertebral level

SpinOUSprOCeSSlength - -. . " Females (mean-SD) -Femates(mean)Males (mean) 40 E úMales (mean+SD) - ¿f____ 35 ---\ 30 ./ Ezs

20

15

10 c3 c7 T1 T6 T10 L1 L5 vertebral level

Figure 15 (cont.): Variables by vertebral levels with mean for males and females and maximum one standard deviation range (male mean+SD, female mean-sD)

F. J. Rühli - Osteometric Variation of the Human Spine 147

Left caudal intervertebrat foramen width

16 Females (mean) 15 -' -' -' Females (mean-So) /---rl-\ 14 - (mean) 13 (moan+sD) -Males 12 -Males- ¿

ErÈ / 10 I I 7 6 L1 L5 c3 c7 T1 T6 T10 vertebral level

Right caudal intervertebral foramen width

16 Females (mean) '- -'-' Females (mean-SD) 15 - -\ ---\ 14 (mean) 13 (mean+sD) -Males- -Males 12 -/ Er 10 9 I 7 6 L5 c3 c7 T1 T6 T10 L1 vertebral level

for males and females and 15 (cont.): variables by vertebral levels with mean Figure female marimum one standarr deviation range (male mean*sD, mean-SD)

t49 F. J. Rühli - Osteometric l/ariation of the Human Spine Sexual dimorphism

As already mentioned above, for the vast majority of the explored variables of the whole sample, males show bigger values than females.

Female mean values were compared as percentages of male ones, with the latter

ones assumed as being I00%. Femur length was on average TYobigget in males, while

femur circumference was approximately llYo different. Similar sexual dimorphism

pattern can be found for the two variables of the humerus. Furthermore, femoral head

breadth shows a sex difference of almost l2Yo. Females present in relation to femur

length a larger bi-iliac width, which is on average just 4Yo smaller than in males.

Females have absolutely bigger values for a large number of intervertebral

foramen widths. These are the only variables examined, of which some are absolutely

larger in females than males. Values absolutely bigger in males, but relative to

percentage difference of femur length de facto larger in females, are additional

intervertebral foramina and a lot of the neural canal measurements, especially the

sagittal dimensions.

In relation to femur length differences, larger values can be found in males,

especially for most of the spinous process lengths, as well as frequently for the pedicle

heights. Furthermore, some values of sagittal and transverse vertebral body dimensions

are also, relative to femur length sex differences, bigger in males. The foramen magnum

dimensions are larger in males than in females, but the sexual dimorphism is for both

diameters smaller than the average femur length sex-difference.

Significant sex differences in mean values, after application of Bonferroni's

correction, were found among the modern samples with proven individuals' sex. For

F, J. Rùhli - Osteometric Variation of the Human Spine 150 most vertebral body dimensions such as height and diameters, for most transverse process widths as well as for pedicle height, there is a signifrcant sexual dimorphism with males showing larger dimensions; whereas for the vast majority of the spinal canal

diameters and intervertebral fårurn"n widths, there is no signif,rcant difference in mean

value between sexes. A complete list of all percentage- and t-values of sexual

dimorphism could be found in appendix 4.

Side dffirences of spinal measurements

possible side difference was tested for the mean values of the bilaterally

measured spinal dimensions, which are pedicle height and intervertebral foramen

widths, in the modern samples. No signifrcant side differences, for both males and

females, have been found for these measures. The t-values, which are non-significant for

any measurement at level p<0.05, even before the application of Bonferroni's correction

for multiple comparisons, could be found in appendix 5'

Inter-correlations of all measurements

The correlations of the osteometric variables with each other show consistent

patterns, which are similar in both sexes. The complete list of all inter-correlations could

be found in appendix 9.

In general, comparable measurements of anatomically closer located vertebral

levels tend to correlate to a higher degree with each other than the same ones located

fuither apart. Additionally, even unrelated measurements, but still closely located in

terms of neighbouring vertebral levels, correlate signifrcantly with each other.

Furthermore, similar measurements even in largely fat apart locations correlate well

151 F. J. Rühti - Osteometric Variation of the Human Spine with each other. There is also a high correlation between the same measurements on both right and left side, as performed here in the cases of the pedicle heights and the intervertebral foramen widths. Tlrpical examples of high vertebral inter-correlations, with a Pearson correlation eoefficient of usually at least approximately 0.6, are ventral versus dorsal vertebral body height, sagittal versus transverse vertebral body dimensions or transverse versus sagittal spinal canal diameters, as measured on the same vertebral level.

The foramen magnum shows primarily significant correlations between its sagittal dimension and the examined sagittal dimensions of the spinal canal. The long bone measurements demonstrate mostly high correlations with each other. Both, femur and humerus show a large number of medium level correlations, but still signif,rcant, with various vertebral measurements. The bi-iliac width shows fewer correlations than other non-vertebral measurements with the vertebral dimensions, but still it expresses a few mild ones, especially with the sagittal vertebral body dimension and the transverse spinal canal dimensions.

Correlation of examined variables with individtml age

The correlation of individual age with the selected spinal and long bone measurements has been tested on the two modern samples; see also Figure 16.

In males, after application of Bonferroni's correction, multiple variables show significant alterations in relation to individual age. At most levels, the sagittal diameter of the vertebral bodies and its transverse diameter show an increase with individual age.

Additionally, the pedicle height shows an increase in size with age. This effect is more clearly visible on the right side than on the left, in the latter one the signihcance

F. J. Rühli - Osteometric Variation of the Human Spine 752 dimensions of the vértebral bodies of the cranial half of the spine and femur

circumference increase then with individual age.

Selected scattergrams of spinal and long bone measurements, significantly

changing with individual age, are presented in Figure 16. The compiete data set on

correlation between the osteometric measurements and individual age at death or major

age groups, respectively, could be found in the appendices 6 - 8'

If one divides the sample not only in the two sexes but also additionally into the

three major time groups and then analyses the correlation between the measurements

and individual age group, the trends found become weaker and less consistent, even

within the same sex.

154 F. J. Rühli - Osteometric Variation of the Human Spine C7 sagittal diameter vertebral body (males)

22 a a

21 a a a 20 ô 't9 aa o Ò a a Et. a a aa o a 17 a a a a 16 a y=0.0602x+ 14'593 a Br = o 3927 a a 15 a a

14 80 50 ô0 70 20 30 40 age (years)

(males) T10 transverse dlameter vertebral body

42 a a 40 I a a 38 a a a a ¡a a t a 36 a a a a a 34 a a E a E a 32 a aa a 30 y=O.lO35x+29 181 H¿ = o.go83 28 a a

26 a

24 80 50 60 70 20 30 40 age (years)

known individual age in Figure 16: Selected variables with siguifrcant change with modern samPles

155 F. J. Rühti - Osteometric Variation of the Human Spine Ll right pedicle height (males)

20 a a

19 a a a a a 18 ', a a a a 17 a a a O a a a '16 l¡' a a E a Ëls a a a 14 a

13 a y=0.0431x+14.167 F¡ 0.2208 12 a =

11

10 60 70 80 20 30 40 50 .age (years)

Minimal humerus c¡rcumference (males)

80 a a a a a 75 a i) a a a a 70 a a aa a a a a a a a a aÒ Ë65 a a a 60 a a a a a a y = 0.1 595x + 56.931 55 A2 =0.2751

50 20 30 40 50 ÞU 70 80 age (years)

individual age Figure 16 (cont.) Selected variables with signifrcant change with known in modern samples

156 F. J. Rühli - Osteometric l/ariation of the Human Spine Mid-femur circumference (males)

110

105 a a a 100 a a a a 95 a a a a a t É ait a a Ës0 a a ôo a o a oÒ 85 a a a a 80 a y=0.1611x+81.035 a a R¿ = 0.1771 75 a 70 20 30 40 50 60 70 80 age (years)

Maximum femur length (females) 490

aa 470 a

a 450 a y=-0.6151x+455.99 R2 =0.2248 a aa a a ! +eo a a ô a a 410 a a a

390 a a

370 25 35 45 55 75 85 age (years)

Figure 16 (cont.): Selected variables with signifrcant change with known individual age in modern samples

t57 F. J. Rühli - Osteometric Variation of the Human Spine Microevolutionary trends since the Late Pleistocene

Ail samples, including the single individuals from the Paleolithic and

Mesolithic, were included to test for signifrcant microevolutionary trends in spinal and long bone osteometry. The i.gr.r.ion models with the highest significance, after application of Bonferroni's correction for multiple comparisons, for each of the examined variables are listed sex-matched in appendix 10. Selected scattergrams of

significant trends are shown in Figures 17 and 18.

In males, with the single exception of the transverse diameter of the vertebral

body at leve1 C3, all other signif,rcant microevolutionary changes of the examined

variables show an increase since the Late Pleistocene. Most significant alterations are of

logarithmic shape. All measurements show for at least one level a microevolutionary

change, most of them for several levels. The foramen magnum dimensions do not show

a signihcant microevolutionary change. All long bone measurements, the bi-iliac width

and the age groups express a significant temporal increase as well.

ln females, most of the significant microevolutionary alterations are of positive

nature as well. Only a few such as e.g., femur length or several intervertebral foramen

widths, decrease through time. The vast majority of the variables show an increase since

the Late Pleistocene. Most of the variables, which show a significant microevolutionary

alteration, follow a logarithmic pattern. Some of the non-spinal measurements, such as

humerus length or bi-iliac width, increase through time in females as well. Again, as in

males, the foramen magnum does not show a significant alteration. Finally, the age

groups show also a positive microevolutionary trend.

F. J. Rühli - Osteomelric Variation of the Human Spine 158 T6 left caudal intervertebral foramen width (males)

18 y = -0.64631n(x) + 16.402 a a 16 a a Hz = 0.1 997 ¡ .aaÒ 14 a I a Ep O a E I a O cO 10 ':t ¡ 1.. o I o

b 100 10 1 00000 1 0000 1 000 years BP

T10 spinous process length (males)

40 a + 37 35 y = -1 .5208ln(x) '323 t a R2 = 0.2334 a a o 30 aa a t o ta, a Ezs a I a 20 a a 15 o 10 100 10 1 00000 1 0000 1 000 years BP

L1 vertebral body sagittal diameter (males) 45 y = -0.6081 Ln(x) + 35.893 R2 0.0798 40 = a o lt 35 a , E a Òa E I a a 30 a a a a a t 25

20 100 10 1 00000 10000 1 000 years BP

Figure 17: Selected variables with signihcant microevolutionary trends in males

159 F. J. Rühti - Osteometric Variation of the Human Spine L1 spinal canal sagittal diameter{males) 24 a a 22 y = -0.3803ln(x) + 19.529 oa a a R2 = 0.0599 o o a 20 a oO l' 3 ö 18 o a E a a E 16 t a a rl 14 o a öa a 12

10

1 00000 1 0000 1 000 100 10 years BP

Maximum femur length (males) 550 530 y = -0.0022x+ 459.97 a 510 R2 = 0.0508 a a 490 a 470 a , ¿so ÈF a a a 430 to I o a 410 a O t a 390 o a a 370 350

1 00000 1 0000 1 000 100 10 years BP

Mid-femur circumference (males) 110 y={.0005x+90.45 a 105 Rz = 0.0398 o a a 100 a o a a a 95 .L a a 'f Eso o aa a È ttO 85 o o o 80 a t t o (Þ a I a o a 75 a a a 70 '100 1 00000 1 0000 1 000 10 years BP

Figure 17 (cont.): Selected variables with significant microevolutionary trends in males

F. J. Rühli - Osteometric Variation of the Human Spine 160 C3 spinal canaltransverse diameter (females) 30 a 28 23.684e'rE-os' Y = I Fì2 = 0.1 85 26 o a a a ËEz¿ a a 22 aa o 20 o o 18 100 10 1 00000 1 0000 1 000 years BP

C7 left caudal intervertebralforamen w¡dth (females) 13

12 y = -0.0001x + 9.8731 t o¡. R2 = 0.0893 11 a I .tt o a aO 10 aa a a. Eo a È a a 8 a I t a a a o I o o a 5 ''1000 '100 10 1 00000 1 0000 years BP

T1 transverse process width (females) 90 o 85 y = -1.1026ln(x) + 78.369 H2 = 0.0832 80 aa t aa 75 a o a Ezo fo oi È a I a O¡, a a 65 a o aa ¡ þU t a 55

50 10 '100000 1 0000 1 000 100 years BP

Figure 18: Selected variables with significant microevolutionary trends in females

F. J. Rühli - Osteometric l/øriation of the Human Spine 761

Secular changes of the intervertebralforamen in the modern samples

The intervertebral foramen was further assessed by linear regression in the modern samples; see also Tables 8 and 9.

A positive secular trend of the slopes for nearly all selected levels of the

maximum intervertebral foramen width, with females demonstrating mostly a stronger

tendency, can be found. For females, on C3, left side only (r:0.77) and bilateral on Ll

(rnro,:0.60, r,"n:0.61), the increase was signif,rcant, even after application of Bonferroni's

correction for multiple comparisons. Other positive secular slope trends, signihcant only

before application of Bonferroni's correction, were found in females on C7 bilateral

(rnr*:0.48, r,"o:0.45), Thl bilateral (r nro¡0.39, r,";0.52), Th6 right (r0.aQ and in males

onCT right (r0.37), Thl bilateral (rnro,:0.46, r,.o:0.33) and L5 left (10.42).

Intervertebral foramen height, as calculated by subtracting pedicle height from

posterior vertebral body height, showed mostly a mild negative secular trend in either

sex, only significant before Bonferroni's correction, in females for C7 bilateral (rnroi-

0.40, q"o:-0.42) aú in males for C7 on the right side only (r-0.35). Intervertebral

foramen heights on Th10 in females and Th10, Ll and L5, all on both sides, in males

were the only ones demonstrating a positive, still insignificant, secular trend.

F. J. Rühli - Osteometric Variation of the Human Spine 163 caudal intervertebral foramen Table 8: Pearson correlation coefficients (r) of width with birth year in modern samples (N total=7l, signihcant at for f.ó.ós before* and after** application of Bonferroni's correction multiple comparisons)

r - Males Level / side r - f,'emales

0.24 C3 / left 0.77**

C3lngh| 0.54* 0.19

0.09 C'7 lleft 0.45*

0.37* C7 lright 0.48*

0.33* Thl / left 0.52*

0.46* Thl / right 0.39*

0.t2 Th6 / left 0.06

-0.03 Th6 / right 0.46*

0.24 Th10 / left 0.27

0.19 Th10 / right 0.35

Ll lleft 0.61* * 0.01

Ll lright 0.60** 0.16

0.42* L5 / left 0.24

0.20 L5 / right 0.2r

164 F. J. Rühli - Osteometric Variation of the Human Spine Table 9: Pearson correlation coefficients (r) of intervertebral foramen height with birth year (N total:71, significant at p<0.05 before* and aftert* applicátion of Bonferroni's correction for multiple comparisons)

r - Males Level / side r - Females

C3 lIeft -0.25 -0.15

C3 / rigþt -0.07 -0.13

C7 lleft -0.42* -0.30

C7 lright -0.40* -0.35*

Th1 / left -0.12 -0.07

Thl / right -0.14 -0.08

Th6 / left -0.29 -0.01

Th6 / right -0.36 -0.06

Th10 / left 0.19 0.05

Th10 / rigbt 0.29 0.03

Ll / left -0.16 0.31

Ll / right -0.17 0.29

L5 lleft -0.20 0.02

L5 / risht -0.03 0.14

165 F. J. Rühli- Osteometric Variation of the Human Spine present Analysis of variance: variable means with respect to time before

An analysis of variance (ANOVA) was performed to test for significant influence of historical age. The various dates before present of the samples and individuals were divided into the three major time groups, with the Paleolithic and

Mesolithic individuals neglected. Additionally, the ratios of sagittal divided by

as well as the transverse vertebral body, foramen magnum or spinal canal diameters'

of the robusticity indices of the long bones, were analysed too' A further subdivision

one of the main age gfoups' samples, not only according to supposed sex but also within

and then the application of an ANOVA, with applying Bonferroni's correction,

further explored' expresses much less significant alterations and has not been in A complete data set of these examinations for both sexes can be found in graphic appendix 11. A summarising graph showing the ANOVA results for means

which are form could be seen in Figure 19, with borderline alterations being the ones,

only signifrcant before the application of Bonferroni's correction.

In males, the ANOVA shows, after Bonferroni's correction for multiple

the transverse width of comparisons, a significant increase at most vertebral levels for

and the spinal canal. Furthermore, some levels of sagittal vertebral body diameters all long caudal intervertebral foramen width show an increase as well. Additionally, a positive bone measurements, foramen magnum length and bi-iliac width show the transverse correlation. A signihcant negative aiteration can be found only for

ratios, the majority of the diameter of the vertebral body at level C3. Of the calculated

significant positive vertebral body ratios and the humerus robusticity index show a

correction ale some change. only significant before the application of Bonferroni's

766 F. J. Rühli - Osteometric Variation of the Human Spine levels of pedicle height, additional intervertebral foramen widths and especially selected levels of sagittal spinal canal and vertebral body diameters.

Furthermore, ANOVA was separately applied for the alterations between the three major time groups, Bronze Age I Neolithic, Medieval and modern times, respectively. As expected, some pairs of time groups show significant differences, but since the other pairs of the same variables do not, the overall change in means for this particular variable will not show a signihcant alteration with time at all, or it will just express one before the application of Bonferroni's correction. For example, there is a significant difference in age group mean between time group 1 and time group 3, but overall there is no such significant difference in males by applying ANOVA for this particular variable. The majority of the pairs showing signihcant differences in means are the Neolithic I Bronze Age time group I versus the modern time group 3. On the other hand, there are variables such as in males e.g., the dorsal height of the vertebral body at level C3 or sagittal diameter of the vertebral body at level Th6, which reveal differences between other pairs of time groups or between all of the time groups. In general, the least frequent significant differences can be found between time group 2

and3, with the majority of such alterations to be visible between time groups 1 and 3. In males, both humerus measurements and femoral head breadth show significant differences between all time groups, whereas for the male femur variable, this is different. Only time group I and 2, which are the Neolithic I Bronze Age versus the

Medieval samples, have significantly different femoral values. Furthermore, bi-iliac width in males does only express significant mean differences between time group 1 and time group 3.

F. J. Rühli - Osteometric Variation of the Human Spine L67 In females, similar patterns emerge. The ANOVA shows, after application of

Bonferroni's correction for multiple comparisons, significant difference in terms of time group for most of the transverse diameters of the spinal canal, as well as some of the intervertebral foramen widths. All these trends are of positive nature. Two levels of sagittal vertebral body diameters, Th6 and Th10, also show signihcant differences in females. Some vertebral variables, such as e.g., additional intervertebral foramen widths, or additional single diameters of the vertebral body or spinal canal, are only significant before the application of Bonferroni's correction. Two levels of transverse diameters of the vertebral body, C3 and L5, show a decrease in size, only signif,rcant before the application of Bonferroni's correction. None of the two foramen magnum dimensions expresses a significant alteration. With the exception of minimal humerus circumference, all other non-spinal variables express a significant positive alteration in females. The calculated spinal ratios and indices show just one with a positive

significant trend, Th6 vertebral body dimensions, but also positive significant trends were found for both humeral and femoral robusticity. The foramen magnum dimension

index shows a signif,rcant decrease in females. Some more ratios in females show

significant alterations, only before the application of Bonferroni's correction. The

further investigation of mean female variables, with respect to major time group pair

differences, shows similar trends to males. Again, there are mean differences between

single pairs of the major time groups, which disappear to be signihcant once all three

major time groups are combined. ln addition, most often significant differences can be

found between time groups 1 and 3. Furthermore, the majority of the long bone

measurements show signif,rcant differences between time groups I and 2, and 1 and 3,

but not 2 and 3, respectively.

F. J. Rühli - Osteometric Variation of the Human Spine 168 By comparing the trends in alteration of variable means between sexes, one finds that most of these significant trends are consistent for both sexes. This is in particular true for some intervertebral foramen widths, selected levels of spinal canal transverse

diameters and sagittal diamè'ters of vertebral bodies. Some trends are only significant in

one sex after Bonferroni's correction, but would be significant, without Bonferroni's

adjustment, in the other sex too. More often hends are significant in males only but not

in females, than the opposite. All trends are consistent in their positive or negative

nature between the two sexes, except for some of calculated ratios.

'1.69 F. J. Rühli - OsleometricVaríation of the Human Spine c3 Mean c7

Thl Th6

Th10 L1

,1 A <.;.> v v

aaa oa

L5 [14¿ls ..... - Male bOfdefline Female Female borderline I Decrease

lncrease 0

Figure 19 Signihcant and borderline alterations of mean values between Neolithic / Bronze Age and modern samPles

F. J. Rùhli - Osteomeh"ic Variation of the Human Spine 770 Mean c3 c7

Th1 Th6

Th10 L1

Male ..... L5 - Male bOfdefline <-+¡ Female Female borderline

lncrease Decrease 0 I

Figure 19 (cont.): Significant and borderline alterations of mean values between Neolithic lBronze Age and modern samples

F. J. Rühli - Osteometríc Variation of the Human Spine 177 Analysis of variance: variable standard deviations with respect to time beþre present

The alterations in standard deviations were examined between the three major 3' time groups by Fisher-test, by comparing differences between time group 1 and A A complete set of these analyses can be found for both sexes in appendix 12' summarizing graph showing the signihcant and borderline alterations of the standard

deviations, the latter changes ones only significant before the application of

Bonferroni's correction, could be seen in Figure 20'

In males, after application of Bonferroni's correction for multiple comparisons,

generally, a significant increase of standard deviations for some of the variables was

found. only the left cranial intervertebral foramen width at c3 and the transverse

of standard process width at level L5 show a significant decrease. A significant increase

vertebral body deviations was found for a few measurements, such as e.g., ventral

L 1 . Muitiple height at level C7 or for transverse diameter of the vertebral body at level

as well as levels of sagittal diameter of the vertebral body and of the spinal canal,

alterations standard selected intervertebral foramen widths, show only significant of

deviations before Bonferroni' s correction'

for the age ln females, a signifrcant increase of standard deviations can be found

the group classification. After Bonferroni's correction for multiple comparisons, only

right craniai intervertebral foramen width on level L5 shows a signif,rcant positive

show a increase of standard deviations. Furthermore, all long bone measurements

positive secular trend for the standard deviations in females. Without Bonferroni's

as few, mostly correction, more intervertebral foramen widths at selected levels as well

cervical, spinal measurements express an increase of standard deviations'

772 F. J, Rühti - Osleometric Variation of the Human Spine No variable shows in both sexes, after Bonferroni's correction, a signifrcant alteration of the standard deviations. In general, more variables in males show

signihcant changes in standard deviations, before or after Bonferroni's correction, than

do in females. The ventral vertebral body height on level C7 and the transverse process

width on level L5 show a significant increase or decrease, respectively, after

Bonferroni's adjustment in males, with females showing a signif,rcant change for this

particular structure only before Bonferroni's correction. The significant alterations in

female dimensions e.g., the long bone measurements do not have significant male

counterparts. Both, C3 and C7 dorsal vertebral body heights, show in males and females

significant increases in standard deviations only before the Bonferroni correction.

F. J. Rühli - Osteometric Variation of the Human Spine 173 c3 Standard Deviation c7

Th1 Th6

Th10 L1

L5 JVI¿ls ..... - Male borderline Female Female borderline I Decrease r lncrease

Figure 20: Signihcant and borderline alterations of standard deviations values between Neolithic I Brot:øe Age and modern samples

174 F. J. Rühti - Osleometríc Variation of the Human Spine Standard Deviation c3- c7

,$

v

Th1 Th6

Th1 0 L1

[vl¿ls L5 -.. .. . Male borderline Female Female borderlíne

Increase Decrease 0 I

Figure 20 (cont.): Significant and borderline alterations of standard deviations values between Neolithic lBronze Age and modern samples

F. J. Rühli - Osteometric Variation of the Human Spine 175 Princþøt components ønølysis of the spinøl aørinbles

Principal component analysis of the spinal measurements ìùras done separately

for each sex and for the-frst five components only. In males, these components

accounted for approximately 49o/o of variation, whereas in females they influence

approximately 57% of the spinal variation. In both sexes, the first components seem to

be linked to size, with the second most influential one to be linked to the size of the

neural pathways. As seen in Figures 21, the two major components, both in males and in

females, do not show a clear trend. A complete data set for the principal components

anaþsis could be found in appendix 14.

F, J. Rühli - Osteometric Variation oJ the Human Spine 176 4

tr 3

tr) 2 tr '6]t -> R É tr c(õ (ú E tr o El o oF <\t 0 E o t! EEt o Þ tr Dú otrE o tr Þ o E (¡t -1 E E o O (ú -2 oÉ I,JJ É. -3 -3 -2 -1 0 2

REGR factor score 1 for analysis 5

Figure 21: Principal components 1 and 2 of males in modern samples

2 E

E E EI Þ tr o P.o o tr EI o -> tr 4 (E ct o (úc O-l o o Àt o b-2o at o () (It -3 o É o tr-4LU -2-10 2 3

REGR factor score 1 for analysis 1

Figure 21 (cont.): Principal components 1 and 2 of females in modern samples

F. J. Rühli - Osteometric l/ariation of the Human Spine 7n I)iscussion

Osteometric htowledge of h-istoric spines

The results of the present study allow a deeper insight into the osteometric variability of the human spine, not only based on sex and individual aging, but also in particular with a special focus on the possible implications of various historic time periods. The osteometric knowledge of historic spines has been elaborated, as could be

seen in Table 1, but, surprisingly, a microevolutionary perspective of historic spines has been mostly neglected so far.

Until now, most measurements of historic human vertebral column had some

limitations either of numerical (small sample size), geographical fiust one major area

covered) or methodological nature (different methods used or just radiological

measurements). Furtherïnore, the majority of previous studies were undertaken with a

direct clinical perspective; see also Table 2. For example, Huizinga et al. (1952) used

19'o century skeletons due to the lack of sufficient recent sources to explore the osseous

dimensions of the lumbar spinal canal with a clinical aim. Scoles et al. (1988) also

mention that the knowledge of vertebral morphology was still limited, therefore, they

provided measurements gained on macerated thoraco-lumbar spine sections'

So far the most similar study on the spinal osteometry from a historic

perspective has been conducted by Jankauskas (1994). He found that the variability of

spinal measurements in historic Lithuanian populations displayed no microevolutionary

trend. According to Jankauskas (1994) the known osteometric spinal data, with their

lack of microevolutionary trends, postulate their restricted value for European inter-

populational studies. Based on the findings of the study presented here, this statement

178 F. J. Rühli - Osteomelric Variation of the Human Spine must be revised at least for some of the spinal osteometry. Jankauskas (1992),

furthermore, did also not report any secular change in the occurrence of spinal

pathologies. This was not an jssue for the present study, but offers a glimpse of how

further research could continue, by focusing on the microevolutionary trends of

particular spinal pathologies as rarely done so far (Rothschild and Rothschild, 1996;

Henneberg and Henneberg, 1999).

To summarize, it is striking to see that historic studies on large spinal samples

and addressing morphometric variations are still rare, whereas for other main body

parts, such studies have been conducted in abundant form and major secular trends a¡e

well known, as already highlighted above. The outline of the study presented here was

to address this lack of knowledge by evaluating the impact of sex, individual age and

historic time period on the morphometry of the human spine in Central Europe; this

despite the awareness of a plethora of possible biases, which are unfortunately

inevitable in such a historic skeletal study.

Study limitations

Microevolutionary changes reconstructed from sometimes very incomplete fossil

and skeletal records are full of pitfalls, such as differences between methods of weight

and stature estimation or completeness of skeleton (De Miguel and Henneberg, 1999)'

The general osteological paradoxes that skeletons in fact represent the non-survivors in

a certain population (Wood et al., 1992), have to be remembered while doing

microevolutionary data interpretation as well. Osteological collections of historic

populations may have an additional selection bias, since some specimens with highest

quality preservation or the ones showing interesting pathologies, which might be

F. J. Rühli - Osteometríc Varialion of the Human Spine 179 completely unlinked to the spine, could have been stored separately. Furthermore, the least preserved skeletons may not be included in any survey at all. For example, in a

10'-12* century cemetery, on\y 82o/o of all vertebrae were preserved (Swedborg, 1974).

These missing individuals might already have been in their lifetime the ones with the most gracile skeleton. Furthermore, usually a large number of the preserved skeletons

show at least macroscopically detectable pathologies, not to mention the ones, which

might have microscopic level alterations making them not to be representative for the

normative healthy population. The ones with macroscopic defects at least were excluded

in a study.

The variability of origin of the selected samples in the present study is another

problem to be addressed. The cultural and geographical-genetical variation of the

included samples might be a possible drawback for a generalization of the findings.

Theoretically, such a study on changing morphology might show results that are more

obvious by focusing on samples from a single location only, by avoiding influences

such as major genetic polymorphism or different environment. Allbrook (1955) already

stated medically important as weil as unimportant variants of the spine could be

resulting from genetical polymorphism. For example, wetzel (1910) in his report on

spinal osteometry highlighted the fact that the European inhabitants differ remarkably.

However, even if there is a high morphological variation present, this may not be true

for all parts of the human body. As Formicola and Franceschi (1996) reported on the

estimated Neolithic body height in Europe, such a variability must not impact on

individual height. They found low standard deviations of less than 4o/o for the total

length of the vertebral column in a vast sample. It is not clear if in the present study the

selected samples of different origin and, therefore, possibly morphological variability,

180 F. J. Rühti - Osteometric Variation of the Human Spine To explore this' . significantly cioud the examined underþing morphological alterations.

sample and would have one must remeasure in a similar way an even mole homogenous to compare the found range-ofvariability'

population, may be The fact that the here chosen sample could represent a biased and especially the highlighted by Martin (1923) saying, that the swiss Neolithic

marks' a Bajuwar sample, show exceptionally strong development of humeral muscle mostly direct sign of individual muscular activity. Therefore, the here selected sampies' the findings that would consisting of South German and Swiss populations, may bias

trends in otherwise be even more obvious. Unlike earlier reports of microevolutionary

people, seem not to show Central European modern Homo sapiens, especially the Swiss

Periods ('wurm, 1982)' In the a major shift in body size at least since the Late Roman from present study especially the tall stature of the Medieval Age samples, originating (1982) explains similar Switzerland and Southern Germany, is astonishing. Wurm

people originating findings with a possible higher content of milk proteins in diet of

through most time of the from the Alpine and swiss area. According to him, in the Alps

between levels modem history intensive stock farming was always present. The trends

would be correct for most of protein intake and adult stature, as shown by Wurm (1982), swiss area' Additionaliy' of Germany, but apparently not for the even more aþine and Wurm (1982) socio-economic factors may interfere with individual stature,

not have been any such impact concludes, that for the highest social classes there might

in stature did not find a on stature at all. Nevertheless, other reports on secular trends

(Henneberg a¡rd Van den Berg' 1990; strong dependence of it on socio-economic levels so, which way social Henneberg, 2001b). Therefore, it is unclear whether and if in

181 F. J. Rühli - Osteomelric Varíation of the Human Spine discrepancies between today geographically Swiss and German populations increased the suggested nutrition-based stature differences.

Additionally, aîy post-mortem alterations of the spinal column are affecting its

morphology. How much the process of skeletonizing alters the spinal morphology has

still to be fully explored. Any macerated bone does not precisely represent its size in

vívo. For the femur, as an example, Martin and Saller (1957) list a post mortem

shrinking of 2.3 mm - 2.6 mrrr, and for the humerus one of 1.3 mm. By including the

cartilaginous part, this amount increases up to 7.1 mm for the femur and 4.1. mm for the

humerus. Todd and Pyle (1928b) addressed the post mortem alterations of spinal

morphology and provide absolute values for the intervertebral discs. Post mortem

alterations of the spine have been discussed in the literature also in particular for the

intervertebral disc (Jacobi, 1927; Adams et al., 1994). The lack of intervertebral disc

and other soft tissue components such as ligamentum flavum cannot be overcome in

osteometric studies. This is in particular true as for the study presented here, if one tries

to establish links between found osteometric alterations and possible clinical symptoms

usually crucially depending on soft-tissue processes. The effect of drying on the

vertebral column seems to reach its final stage after a few weeks and contributes to a bit

less than 3o/o of the total column length, which if far more than for other human bones

(Todd and Pyle, 1928a). Furthermore, the extent of drying of the vertebral column

seems to vary for all parts at least of the vertebral body. Todd and Pyle (1928a) found a

lower relative shrinkage for the ventral aspects of the vertebral body and declare any

shrinkage of the articular processes to be negliable. Therefore, osteometric measures do

always slightly differ fromin vivo dimensions.

F. J. Rühli - Osteometric Variation of the Human Spine 182 Furthermore, intra vitam pathologies affect the spinal morphology. Minor osteophytic alterations were not a reason for exclusion of vertebral columns from the present study as long as they were regarded as conìmon age-related adaptations and did not interfere with the selected measurements. To highlight this, one has to be aware of the high frequency of such alterations as already reported earlier (Bailey and Casamajor,

191 l; Hurxthal, 1968; Hukuda et a\.,2000). For example, Nathan (1962) stated that in a sample of 400 recent vertebral columns by un age in the forties all individuals showed at least early stage osteophytes on some of the vertebrae and, therefore, such mild changes can not be regarded as a pathology. Jankauskas (1992) found the onset of spinal degenerative changes to be in his archaeologic sample at around 25-30 years of age for osteophles or even younger in cases of Schmorl's nodes. In studies of cadavers of various inter-populational origin, done by Eisenstein (1977; 1980), between 25 o/o and

56Yo of the skeletons showed some form of osteophfes. Also Park (1980) lists that 95% of people aged 70 years of both sexes will show age-related degenerative spondylosis in the lower lumbar spine. In a clinical study involving individuals who did not show any neurological signs, Pallis et al. (1954) found on X-rays of the cervical spine in a sample after 50 years moderate or severe canal narrowing or foraminal narrowing in76Yo and

72Yo respectively. Surprisingly, the prevalence and severity did not further increase in this sample after 50 years of age. Marginal osteophytes at the ventral border of the vertebral bodies were present n82o/o of individuals. Based on all these reports, one may more easily approve the chosen approach in terms of minor age-related spinal alterations. Secondary degenerative changes, which involve osteophytes or soft tissue alteration, such as increased thickness of ligamentum flavum or bulging of the

F. J. Rühli - Osteometric Variation of the Human Spine 183 intervertebral disc, are the main etiologies of most cases of spinal stenosis in modem clinical situations. These changes caí,not be explored in such microevolutionary osteometric study including non-degenerative spinal columns only. Nevertheless, there are significant correlations between the osseous and soft tissue aspects of the spinal column described, such as between the posterior disc and intervertebral foramen height or between the cross-sectional areas of the foramen and the related nerve roots

(Hasegawa et a\.,1995). Thus, by obtaining osseous measures, to a limited extent only one may assess the living soft tissue involving conditions.

How far the osseous outline of the spinal canal and its major content, the spinal cord, are correlated, needs to be further evaluated. Preliminary results by Humphreys et al. (1998) show that the ratio of these two structures in the cervical spine changes during adulthood. If there were a consistent conelation of these two structures, this would help to draw conclusion on neural pathways by obtaining osseous measurements only.

The true size of the intervertebral foramen, as another example, can only roughly be assessed by its known osteometric diameters. Even plain radiography does not allow accurately enough to determine this crucially on the presence of soft tissue depending structure(Stephens eta1.,1991).Toassesstheoverallsizeoftheintervertebralforamen it would be necessary to know the height of the intervertebral discs as well as their contribution to the height of the intervertebral foramen. Therefore, one has to rely for this on data gained from clinical or cadaveric studies (Jacobi, 1927; Yu et al', l99l;

Humphreys et a1.,1998; Tribus and Belanger, 2001).

Furthermore, the rather small sample sizes in historic spinal studies have

statistical advantages and disadvantages. Ty'pe I-errors are limited, but the ability to hnd

F. J. Rühli - Osteometric Variation of the Human Spine 784 ' real findings is more diffrcuit, resulting in type II errors. The critical sample size, the extent, and the importance of possible errors of measurement have already been

addressed above. . i

Beside genetic influences or individu al age, clinical conditions, such as fractures,

drug application or various bone diseases, influence the spinal morphometry' Without

background information on historic skeletons, it may be hard to know if such an altering

situation was present and the gained data can be regarded as normative at least for the

time period and geographic background only.

Another issue is raised by the question how far osteometric findings on a

particuiar vertebral level can be generalized for neighbouring levels or whole spinal

regions. To address the interrelation between osteometric spinal measurements

inter- Jankauskas (1994) performed a cluster-analysis with both sexes pooled, since

sexual differences in correlation coefficient were minimal. He found two main clusters:

one of longitudinal measurements and one of the transverse diameters. The inter-sexual

present differences are not negligible in the study presented here. Nevertheless, in the

study dimensions of vertebrae are most strongly correlated with each other at

neighbouring levels, as already found in earlier studies (Hermann et al',1993), as could

also seen in Table 10. As also listed in appendix 9, similar measurements of different

vertebral levels correlate generally better than non-related measurements. To conclude,

present study' one may based on the data provided by Herrm ar:rr-, et at. (1993) and by the

mostly assume that by comparing selected vertebral levels a found trend can be

generalized for the whole vertebral column.

185 F. J. Rühli - Osteometric Variation of the Human Spíne levels, Table 10: Inter-correlation of anterior vertebral body height at various measured on X-rays; *:signif,rcant at p<0.05 (Hermann et øl',1993)

Level Males Females

(N:43) (N:70)

Th6 L1 L5 Th6 L1 L5

Th10 0.29 0.56* 0.29 0.52* 0.45* 0.45*

LI 0.28 0.53* 0.35* 0.45*

L5 0.23 0.29 0.51* 0.45*

186 F. J. Rühli - Osteometric Variation of the Human Spine Comparative analysis of the results

It has been assumed and shown in previous work that spinal morphometric ratios follow a normal or Gaussial distribution (Minne et aL.,1988; Black et a|.,1991;Xu et at., 1995). This is the case, for most of the investigated spinal traits in the present study as well. Similar sample sizes for both sexes were chosen and both sexes show overall similar age distribution facilitating further the interpretation of the results.

The main vertebral body diameters were measured in the present study since they reflect major mechanical players of the spine, as already outlined above. Piontek

(1973) found an increase of massiveness of the vertebral bodies caudally' This seems to be related to the increased load bearing. It is well known that such loading on the spine can be much higher, depending on the body position, than only the normally neutral up to 60Yo of total body weight in the lower lumbar spine. Silva e/ al. (1997) declared that trabecular anisotropy of the human bone is crucial in load distribution within the spinal

column. Thus, it would be worth further investigation how trabecular anisotropy not

only changes within individuals, but also if it shows any detectable microevolutionary

trend. The reports of vertebral body dimensions with measurements comparable to the

ones used in the present study, with means for the whole sample as well as the mean for

the modern subgroups, are listed in Tables 11-15 and 17. One can see that the vast

majority of the osteometric dimensions measured in the present study fall clearly within

the range of earlier reports. Furthermore, one may notice the wide range of reported

osteometric values, which might have been caused by the geographically and stature-

wise heterogenic samples. This will be further addressed below, with a particular focus

on the influence of individual stature on vertebral dimensions for the present study.

F. J. Rühli - Osteometric Variation of the Human Spine 187 Table 11 Vertebral body height (mm) of various samples; measurements similar to Martin (1928)

Level / Sample Ventral Dorsal Ventral Dorsal Reference males males ales c3

Stewart (1962) Shanidar I 1 1,0 12.5

Ditto Skhul 1 10.5

Heim (1976) La Ferrassie 1 t2.5

Matiegka Predmosti 3 l1.3 t2.6

Ditto Predmosti 14 13.0 13.0

Ditto Predmosti 4 10.4 13.0

Ditto Predmosti 10 I 1.4

Jankauskas (1994) Lithuanian Paleopopulations - 1" / 2"0 13.4 13.9 t2.4 12.5 Millenium AD (N males=159, 160; N females=109, I l3)

(1973) Early Medieval Polish (N males:48, N 13.3 12.0 Piontek females:25)

(1966) Polish 12th centurY (N=l) t2 t4 Kaliszewska

Piontek and Budzynska (1972) Rural 12*-14'o centurY Polish 13.4 15.2 12.7 14.3 (N males=19, N females=16)

Piontek and Zaborowski (1973) l2'o-18'n century Polish (N rnales=25, N t4.3 13.8 12.5 t2.7 females=25)

13.9 Ditto Urban 14'o-18'o century Polish (N males=l8, 14.3 15.2 12.5 N females=I4)

Kandziora et al. (2001) Germans (N males=l0, N females=I0, both 15.3 15.4 sexes combined)

t2.5 14.0 tl.2 13.6 Taflinska, cited by Piontek and Polish Q.{ males=56, N females=44) Budzynska

Hasebe (1913) Japanese (N males=20, N females=I0) 14.4 t5.2 12.8 t4.2

cited by Piontek and Budzynska Bushmen (N males=24, N females=l5) I 1.4 t3;7 10.8 r 3.3 Duparc,

Knrczkiewicz, cited by Piontek and Australians (N males=16, N females=I0) 1 1.5 t4.2 10.0 13.3 Budzynska

Cyriax (1920) English (?, N both sexes = +/-'10) 12.9

Tominaga et al. (1995) Recent Americans (?, N males=4, N females= 13.8 t4.2 2, both sexes combined)

Aeby (1879) Recent Europeans (N=8) 11.9 l2.l

188 F. J. Rühti - Osteometric Variation of the Human Spine (1883) Recent Europeans (N=?, both sexes?) 12.4 Andenon

(1939) American Whites (N=+/-96) 14.1 14.0 Lanier

Modern French (N=?) 12.5 Ditto (l99la) Recent Americans (N = 12, both sexes) - I l 1.6 Panjabiela/.

Thomson (1913) Recent Europeans (N=3,6o,¡ t"*.., 13.7 l3

Ditto Recent Bushman (N males=l, N females=l) 9 l0 10 10.5

Stewart Range of global sample (N=20) l1 - 17 10-15

12.4 Present study (whole samPle) f3.7 14.0 t2.3

Present study (modern subgroups) l4.l 74.6 12.5 12.8

c7

shanidar I t¡.0 14'o Stewart

Shanidar 2 13.0 14'0 Ditto

La Chapelle-aux-saints f 10.6 Trinkaus (1985)

La Chapelle-aux-saints I 13.4 Heim

Predmosti 3 13.0 14.5 Matiegka

Predmosti 9 13.0 14 0 Ditto

predmosti 14 16.0 Ditto

Predrnosti 4 I 1.0 12.3 Ditto

Dolni Vestonice 15 t4;7 15.3 Trinkaus þers. comm.)

Lithuanian Paleopopulations (N males=l72, 14,1 15.0 t3.2 13.8 Jankauskas 183; N females=l 18, 126)

Early Medieval Polish (N males=50, N 14.1 13.5 Piontek females= 32)

Polish 12th century 14 14 Kaliszewska

Piontek and Bud4mska Rural 12"-14'o century Polish 14.2 18.6 13.8 t7.6

16.6 Ditto Urban 14'o-18'o century Polish 14.2 17.6 t4.2

Piontek and Zaborowski 12t-18ú century Polish I4.2 15.6 t4.2 t4.3

Kandzioø et al. Germans (both sexes) 15.1 l5'3

Polish 13.2 16.8 13.0 15.3 Taflinska

Japanese 14.5 16.8 13.4 15.3 Hasebe Bushmen 12.6 l4'8 t2.2 t4.2 Duparc

Australians 12.6 l4'9 I 1.8 13.8 Kruczkiewicz

189 F. J. Rühti - Osteometric Variation of the Human Spine English (?, both sexes) 13.4 Cyriax

s¡ al Recent Americans (?, N males=4, N females= 15.8 16. I Tsni¡¡g¿ 2, both sexes combined)

Aeby Recent Europeans 12.3 13.3 - i Recent Europeans (both sexes?) 13.0 A¡denon

Arnerican Whites 14.4 15.0 Lanier

Modern French 13.0 Ditto

Thomson Recent Europeans þoth sexes) 13.5 t4

Panjabi et al. Recent Americans þoth sexes) t2.8

I 1.5 Ditto Recent Bushman I I 10.5 11

Range global samPle 11.5 - 12 - 16.5 Ste$/art 16.5

Present study (whole sâmple) 13'9 14.9 t2.9 13.6

Present study (modern subgroups) l3'7 1s.3 12.8 13.6

Thl

La Chapelle-aux-Saints I 14.0 Heim

Matiegka Predmosti 3 t7.5

Ditto Predmosti 9 15.0

Predmosti 14 17.0 18.0 Ditto

Ditto Predmosti 4 14.8 15.6

Dolni Vestonice l5 15.2 Tri¡kaus

Lithuanian PaleopoPulations 16,0 17.4 15.1 16 I Jankauskas (N males=169, 184; N females=l 15, 126)

Early Medieval Polish (N males:48, N 16.4 15.4 Piontek females= 38)

Polish 12th centurY t6 t7 Kaliszewska

Rural l2'-14' century Polish t6.9 19.6 16. I 17.8 Piontek and Budzynska

Ditto Urban 14"-18'o centurY Polish 16.5 18. 1 t4.2 r'7.1

Polish ts;t 18.2 t4.9 15.2 Taflinska

Cyriax English (?, both sexes) 15.5

Japanese 15.7 16.8 14.8 l5,4 Hasebe

Bushmen 14.4 t4.9 13.4 13,8 Duparc

Australians 14.5 14.5 13. I 14. r Kruczkiewicz

L90 F. J. Rühli - Osteomelric Variation of the Human Spine Jacobi (1927) Recent Gerrnans (N=102, both sexes) r5.2 I5.7

15.3 Aeby Recent EuroPeans 14.3

15.9 Anderson Recent Europeans @oth sexes?) 14.8

.16.2 Lanier American Whites t't.3

Todd and Pyle (1928b) American \ilhites (N=43) 15.9 t7.7

Ditto Modern French 14.5

Thomson Recent EuroPeans (both sexes) 15.5 16

Panjabi ef a/. (199lb) Recent Americans þoth sexes) 14.1

13.s Ditto Recent Bushman t2 I 1.5 l3

t4.6 15.6 Present studY (whole samPle) 16.0 r7.2

t4.s 15.7 Present studY (modern subgrouPs) 16.0 t1.3

Th6

t7.3 Matiegka Predmosti 4

19.6 Jankauskas Lithuanian Paleopopulations l9'l 2r.0 17.8 (N males= 152, l'70 N females= 103' 108)

Piontek Early Medieval Polish (N males=5O, N l9'8 18.3 females:41)

Kaliszewska Polish 12th century 19 l9

Piontek and Budzynska Rural l2'-14'o centurY Polish 20.1 2't;7 t9.2 23.6

22.7 Ditto Urban 14'o-18" century Polish 20.0 26.0 18.5

Marchesi ef al. (1988) Swiss (N males= 18, N females:l5, both sexes 18'6 20.8 and sides combined)

Cotterill er a/. (l 986) Canadians (recent?, N:10, both sexes?) 17 '5

Cyriax English (?, both sexes) 18'3

polish 18.9 26.2 18.0 23.6 Taflinska

Japanese l9'0 23.9 17.0 21.2 Hasebe

Duparc Bushrnen 17.3 20.4 t6.4 20.0

Australians 16'6 2t.9 15.1 t9.4 Kruczkiewicz

Jacobi Recent Germans (both sexes) l'l 'l 19.0

Parrjabi et al. Recent Americans (both sexes) 17.4

19.5 Aeby Recent EuroPeans 16.9

Anderson Recent Europeans (both sexes?) 18'l 19.9

791. F. J. Rühti - Osteometric Variation of the Human Spine Lanier American lYhites 19.0 20.8

Todd and Pyle American Whites 18.7 20.6

Thomson Recent Europeans þoth sexes) l9 20.7

I r6.5 Ditto Bushman l'l l8 t7

17.5 t9.2 Present studY (whole sample) 19.0 20.9

17.7 19.8 Present study (modern subgrouPs) 19.0 21.0

Th10

Matiegka Predmosti 3 2t.9 2t.9

Ditto Predmosti l4 22.4

18.2 Ditto Predmosti 4 t't.5

22.4 Ditto Predmosti 10

Trinkaus Dolni Vestonice 15 23.2

Jankauskas Lithuanian PaleoPoPulations 2t.4 23.6 20.3 21.9 (N males=l52,, 160; N females= 95' l0l)

Kaliszewska Polish, 12th centurY 2l 22

11 ',' 27.6 Piontek and Budzynska Rural 12"-14'o centurY Polish 3 1.9 20.9

27.! Ditto Urban 14"-18" centurY Polish 22.9 31.3 2t.9

))1 Rosenberg (1899) Late 196 centurY Dutch (N=3' sex?)

26.9 Taflinska Polish 2t.8 28.9 20.9

19.2 24.5 Hasebe Japanese 21.5 28.2

Marchesi el a/. Swiss (both sexes and sides combined) 2t.l 23.2

Cyriax English (?, both sexes) 21.2

23.3 Duparc Bushmen 19.9 24.6 18.5

1)) K¡uczkiewicz Àustralians 20.1 24.5 18.6

Jacobi Recent Germans (both sexes) 21.0 2r.l

22.2 Aeby Recent Europeans 21.7

Andenon Recent EuroPeans þoth sexes?) 2r.0 22.9

22.3 Thomson Recent Europeans @oth sexes) 2l

19 Ditto Recent Bushman (N males=2, N females=l) 20 19.8 19

28.7 Diachti er a/. (1995) Italians, premenoPausal (N=50) 28.2

26.8 Ditto Italians, postmenopausal (N=76) 26.2

192 F. J. Rühli - Osteometríc I/ariation of the Human Spine Recent Americans (both sexes) 20.2 Pafiabi et al.

Àmerican Whites 22.3 23.7 Lanier

American Whites 2t.5 23.1 Todd and Pyle

i'r.r Present study (whole samPle) 7 23.7 20.9 21.7

Present study (modern subgrouPs) 22.2 23.8 21.4 22.1

LI

Predmosti 10 25.0 Matiegka

Dolni Vestonice 15 22.4 26.7 Trinkaus

26'3 23 25.5 Vallois (1977) Téviec Q.,l males=3, N females=4) 23.3

Lithuanian Paleopopulations (N males= l7l, 24.9 2'7.8 24.5 20.4 Jankauskas 180 ; N females= 96, 103)

Early Medieval Polish (N males=50, N 26'3 25.3 Piontek females:45)

Polish 12th century 22 2'7 Kaliszewska

Rural 12'-14* century Polish 26.1 34.4 25.5 29.5 Piontek and Budzynska

Urban 14'o-18'o century Polish 26'4 32.8 25.3 288 Ditto

Late 19ü century Dutch (sex?) 24 Rosenberg

English (?, both sexes) 24.4 Cyriax

Polish 24.9 30.4 24.5 29.5 Taflinska

Ma¡chesi ef aL Swiss (both sexes and sides combined) 25.9 2'l .2

Japanese 23'2 25.9 23.8 26.7 Hasebe

Bushmen 22'l 27.3 22.7 24.6 Duparc

Australians 25 '3 )1.4 22.0 25.1 K¡uczkiewicz

Americans (N=30, both sexes) 25.0 25.8 Berry et a/. (1987)

Recent Germans (both sexes) 24,5 25.7 Jacobi

et al. Recent Americans (both sexes) 23.8 Pnjabi

Recent Europeans 25.6 26.0 Aeby

Recent Europeans (both sexes?) 24.6 26-5 Anderson

Italians, premenoPausal 33.1 31.3 DiactnTi et al.

Italians, postmenopausal 29.4 3l.4 Ditto

American Whites 26.2 28.3 Lanier

Pyle American Whites 25'7 2'l '3 Todd and

793 F. J. Rühli - Osteometric Variation of the Human Spine Thomson Recent Europeans @oth sexes) 24.3 26.3 )') Recent Bushman (N males=2, N females=l) 22.3 23.8 2t.5 Ditto

(2001) Recent Europeans (N males =2; N females=2, 28 Boszczyk et al. combined)

Present study (whole samPle) 25.8 28.0 24.7 26.3

Present study (modern subgrouPs) 25.5 27.9 25.0 26.4

L5

Matiegka Predmosti 3 29 '4 23.0

Predmosti 14 20.5 Ditto

Predmosti 4 2'7.0 20.6 Ditto

Predmosti 10 23.5 Ditto

Téviec (N males=4, N females=3) 24'5 20.5 22.2 22.5 Vallois

Lithuanian Paleopopulations (N males= 170, 28.0 23.4 26.2 22.2 Jankauskas 188 ; N females=lOs, 124)

Early Medieval Polish (N males--48, N 29.4 27.7 Pio¡tek females= 41)

Polish 12th century 22 Kaliszewska

Rural 12*-14'o century Polish 29.3 35.7 28.1 32.6 Ditto

Piontek and Budzynska Urban 14'o-18'o century Polish 29.4 34.3 28.2 32.9

Late 19'century Dutch (sex?) 227 Rosenberg "7 English (?, both sexes) 2'7 -8 Cyriax

Potish 28.7 34.7 26.9 32.4 Taflinska

Marchesi e/ ai. Swiss (both sexes and sides) 28.9 24.7

Japanese 27.5 34.6 25.6 3 1.8 Hasebe

Bushmen 24'4 30.s 24.8 30.1 Duparc

Australians 24.3 30.8 23.0 29.9 K¡uczkiewicz

Panjabi et al Recent Americans (both sexes) 22.9

A¡nericans (both sexes) 28'7 23.1 Berry et al.

Recent Europeans 29.8 23.6 Aeby

Anderson Recent Europeans (both sexes?) 27.2 222

Italians, premenoPausal 35.3 32.5 Diacinli et al.

Italians, postmenoPausal 34.1 30.6 Ditto

194 F. J. Rühli - Osteometric Variation of the Human Spine American \ilhites 28.9 29.1 Lanier

American Whites 28.1 23.7 Todd and Pyle

Recent Europeans þoth sexes) 29 2l Thomson

Recent Bushman (N males:2, N females:l) "24.3 22 23 20 Ditto

Recent Europeans þoth sexes) 2f Boszcryk et al.

Present study (whole sample) 28'6 24.5 27.0 23.4

Present study (modern subgroup) 28,9 24.1 28.1 23.6

F. J. Rühli - Osteometric Variation of the Human Spine 195 measurements Table 12: Vertebral body diameters (mm) of various samples, similar to Martin (1928)

Sagittal Transverse Reference Level / SamPle Sagittal Transverse males females

c3

2t.3 Piontek (1973) Early Medieval Polish (N males=48, N 156 233 14.0 females=2í126)

20.0 142 18.6 Piontek and 12'o- l8'o century Polish (N males=25, N 16.0 females=25) (re73)

Cyriax (1920) Engtish (N both sexes=+/-70) 20.9 Aeby (1879) Europeans (N=3, both sexes) t5.2 23.8 Ardenon (1883) Europeans (N:28, both sexes?) 15.2

13.3 2t.0 Thomson (1913) Europeans (N males=5, N females= 8) 15.0 23.1 t2 Ste{ko (1926) Russians (N males=28?, N females=10?) t3 23.s t2 2l Ditto Bushmân (N males=l, N females=l) t2.5 l8

14.8 18.5 Present studY (whole samPle) 16.0 19.3

t4.7 18.1 Present study (modern subgrouPs) t6.2 19.3

C7

16.8 27.5 Piontek Early Medieval Polish (N males=5O, N r8.3 29.4 females=32)

27.0 t6.2 25.6 Piontek and 12'o-18n century Polish t6.9

Zaborowski

15.6 26.2 Aeby Europeans t6.2 285 Andenon Europeans (both sexes?) 18.3

3l .3 Thomson Europeans þoth sexes) l6 Cyriax English (both sexes) 29.2

t4 Stefl

t2 26 Thonson Bushmen (N males=2, N females=l) l4 27

15.6 24.8 Present studY (whole sarnPle) r7.l 26.5

1ó.0 24.4 Present study (modern subgrouPs) t't.7 26.6

196 F. J, Rühti - Osteomehic Variation of the Human Spine

31.0 Present study (modern subgrouPs) 31.3 34.7 27.3

LI

29.6 42 Piontek Early Medieval Polish (N males=5O, N - f3.2 47.2 females=45)

Berry et al. Americans (l'{:30, both sexes) 28.9 39.5

Cyriax English (both sexes) 39.2

Postacchi¡i e¡ ¿/. Italians (N = 63, both sexes) 29.0 41.0 (re83)

Ditto Indians (N=58, both sexes) 25.0 36.0

29.3 4l,3 Aeby Europeans 32.7 46.0 A¡derson Europeans (both sexes?) 29.9

Thomson Europeans 28.7 37;1

28 Stefl

26.7 3 8.8 Scoles af ai, (1988) Americans 29.5 44.3

)q) 26.r Amonoo-Kuofi (1985) Nigerians (N males=79, N females=43)

27 34 Eisenstein (1977) Caucasoid (N males=78, N females=35) 3l 39

25 35 Ditto Zulu Negroid (N males: 108, N females=54) 28 39

t< 34 Ditto Sotho Negorid (N males= 106, N females=62) 27 38

27.6 3s.5 Present studY (whole sarnPle) 31.'l 40.3

28-2 3s.9 Present study (modern subgrouPs) 32.9 41.0

L5

325 50.3 Piontek Early Medieval Polish [N males:48, 35 55.2 females=41/43)

Europeans 36'2 s4.0 31.4 s06 Àeby

Postacchini e/ a/ Italians (both sexes) 33 0 49.0

Ditto Indians (both sexes) 29'0 43.0

Berry et al. Americans (both sexes) 32.4 46.r

e/ ¿/. Americans 34.5 52.9 31.5 48.6 Scoles Cynax English (both sexes) 48.0 Nigerians 34'2 3l .3 Amonoo-Kuofi

36'5 Andenon Europeans @oth sexes?)

198 F. J. Rühti - Osteometric Yariation of the Human Spine Thomson 30.7 42 Europeans @oth sexes)

54 25 Stefl

3 8.5 25 34 Ditto Bushmen (N males=2, N females=l) 29.5

46 30 42 Eisenstein Caucasoid 3f

31 43 Ditto Zulu Negroid 32 45

44 3l 42 Ditto Sotho Negorid JJ

47.8 31.1 44.1 Present studY (whole sarnPle) 33.6

47.7 30.4 42.6 Present studY (modern subgrouPs) 34.5

199 F. J. Rühti - Osteometric Variation of the Human Spine The maximum pedicle height was also explored in the present study, since it could reflect any morphological alterations in particular as a bridging structure between the vertebral body, the laminae and the transverse and spinal process. Pedicle robustness is linked to pedicle function in distribution of force and columnar stress (Shapiro, 1993;

Sanders, 1998). Sanders (1998), who basically divided the human spinal column into just two force bearing pillars, already highlighted the extreme steady demand for the pedicles to support bending stress, due to their physiological positions between the two main force bearing pillars, the frontal vertebral bodies and intervertebral discs and the dorsal pillars, consisting of the laminae and the zygoapophyseal joints. Sanders (1998) also emphasizes the importance of the interaction with the ilio-lumbar ligament as

another factor in developing typical human lower lumbar pedicle size. Therefore, any

alterations of mechanical properties on the human spine would most likely be reflected

on the pedicle size. If the increased pedicle area in humans links to the unique upright

locomotion is still controversially debated (Davis, 1961; Shapiro, 1993) and was not an

issue in the present study. For clinical pu{poses, Banta et al. (1989) recommended to list

rather maximal values instead of the usual standard deviations for reports on the pedicle

size. Nevertheless, the particular effective dimension of the pedicles they measured is

not of real value for osteometric analysis.

The impact of individual stature on pedicle dimensions has been addressed

equivocally so far. Scoles et al. (1988) found no clear link between pedicle dimensions

and individual size, unlike for the correlation between vertebral body height and stature.

On the other hand, Karaikovic et al. (1997) describe a correlation between pedicle

F. J. Rühli - Osteometric Variation of the Human Spine 200 mostly the case in the present dimensions and individual body height, which was also study. with the A tisting of major earlier reports of pedicle height dimensions together

modern sample could be found means of the whole sample presented here as well as the inTable l3

201 F. J. Rühti - Osteometric Variation of the Human Spine Table 13: Maximum pedicle height (mm) of various samples

- Level / Sample Pedicle height - Pedicle height left Reference right

c3 11 Panjabi e/ a/. (I99la) Americans (both sexes, N=12) 7.6

Tominaga et al. (1995) Recent Americans (?, N males=4, N females:2, both 7.5 sexes combined, side?)

Ebraheim et al. (1991) Recent Àmericans (?, N males=25, N females=I5, 6,8 (m) / 4.7 (Ð? both sides combined)

Kandziora et ø1. Germans (N males=10, N females=l0, both sexes 7.4 Q00l) combined, side?)

(m) 6.1 (f) Present study (whole samPle) 6.9 (m) / 6.1 (f) 7.0 /

(m) (Ð Present study (modern subgrouPs) 7,3 (m) / 6.3 (f) 7.3 / 6.2

C7 Panjabi et al. Àmericans (both sexes) 7.5 7.5

Xtt et al. (1995) Americans (N males=32, N females=24, side?) 7.1 (m) / 7'0 (Ð

'lominagr et al. Recent Americans (?, N males=4, N females=2, both 7.4 sexes combined, side?)

Ka¡rdzior¿ et al. Germans (both sexes, sides?) 8'5

(m) (f¡ Present study (whole samPle) 7.2 (m) I 6.6 (f) 7.3 / 6.6 7.s (n) / 6.s (Ð Presents study (modern subgroups) 7.s (n) / 6.6 (Ð

Th1

Panjabi ¿/ a/. (199lb) Americans (both sexes) 9.3 9.9

Scoles et ai. (1988) Americans (N males=25, females=25; side?) e.2 (m) / 8.4 (Ð

(m) 8.a (Ð Present study (whole sample) e.2 (m) / La (Ð e.a /

(m) 8.4 (f) Present study (modern subgrouPs) 9.1 (m) / 8.3 (f) 9.3 /

Th6

Vaccaro et al. (1995) Americans / Asians (N males= 8, N females=9, both 10.1 sexes combined, side?)

202 F. J. Rühti - Osteometric Variation of the Human Spine Arnericans (both sexes) 12.0 I 1.6 Panjabi et al.

Germans (?, N:4, both sexes?) I I.4 Kothe et a/. (1996)

Americans l1.s (m) / 10.6 (Ð Scoles ef a/.

Present study (Ìvhole sample) 12.2 (m) / 10.s (Ð r2.0 (m) / 10.4 (f)

Present study (modern subgroups) 12.6 (m) / 10.8 (Ð 12.2 (m) / 10.s (Ð

Thl0

Aniericans (both sexes) 14.7 15.0 Panjabi et al.

Chinese (N males=25, N female=15, side?) r4.4 (m) / r4.2 (Ð Hota et al. (1993)

Americans / Asians (both sexes, side?) 14. l Yaccaro et al.

Present study (whole sample) 1s.4 (n) / 14.0 (Ð ls.s (m) / 13.9 (f)

Present study (modern subgroups) rs.8 (m) / 14.3 (D rs.7 (m) I 14.3 (Ð

L1

Indians (N males= 18, N females=2; side unknown) 1s.7 (m) t ts.7 (Ð Mitra et al. (2002)

Americans (N males=38, N females=3l; side?) 17.0 (m) / ls.3 (Ð Olsewski et al. (1990)

Chinese (side?) 1s.e (Ð / ls.s (Ð Hott et al.

Americans (both sexes) 15.9 15.8 Panjabi el al. (1992)

Arnericans 15.3 (m) / 14.s (Ð Scoles ef a/.

Americans (N=30, both sexes) 15.6 15.6 Bcrry et al. (198'1)

Present study (whole sample) r6.0 (m) / r4.s (Ð 1s.7 (m) / 14.3 (f)

Present study (modern subgroups) 1ó.s (m) / 14.6 (f) 16.4 (m) I 14.4 (Ð

L5

Indians (side unknown) ls.7 (m) / 17.0 (Ð Mifa et al.

Americans (N males=47, N females=39, side?) 17.4 (m) I 16.2 (Ð Olsewski ¿/ a/.

A¡nericans (both sexes) 180 19.2 Panjabi et al.

Americans 16.2 (m) / l8.s (Ð Scoles e/ ¿/.

F. J. Rühli - Osteometric Variation of the Human Spine 203 Americans 13.8 13.6 B,erry et al.

Hou et al. Chinese (side?) 20.5 (m) / 18.7 (Ð

Present study (whole sample) 14.6 (m) / r3.s (0 1a.0 (m) / r2.8 (Ð

Present study (modern subgrouPs) la,s (m) / 13.3 (Ð 13.9 (m) / 12.7 (f)

F. J. Rühli - Osteometric Varíatíon of the Human Spine 204 The size of the neural canal is a crucial osteometric dimension. The relation between spinal cord and osseous spinal canal size may be important for clinical issues,

suggest as pointed out by Panjabi et at.(l99la) for the cervical spine. For example, they

C6 to a possible link between the decrease of the spinal canal I spinal cord ratio from

C7, andthe subsequent high vulnerability to neural damage, and the high rate of spinal

young cord injuries at this level; as shown by Fife and Kraus (1986). Furthermore, in a asymptomatic clinical sample, Schmid et at. (1999) found no body position dependent changes of the cross-sectional areas of the spinal canal when measured at the osseous level, whereas the same measurement on the disc level did change' This means for the present study that due to its independence of body positions, at least in asymptomatic

individuals, the spinal canal dimensions at the vertebral body levels may be used for

at the comparison between clinical and skeletal samples. The spinal canal dimensions

disc level cannot be assessed in osteometric studies any\,vay.

One has to wonder, how far osseous spinal canal dimensions reflect its content'

By having bigger spinal cords, more muscular individuals, may also need larger bony

predispose neural spaces; unless they show smaller reserve capacities, which then would

them for spinal pathologies. Surprisingly, there is a sexual dimension issue as well.

Female and male mammals have similar size of neural nerve roots, as described by

Dunn (1912),therefore, relative to body weight, female ones are even bigger than males

in this report. ln the case of the cervical nerve root, as examined by Dunn (1912), this pelvic cannot be due to a higher sex dependent visceral demand such as e.g., in the

region, but must be linked to a higher periphery somato-motoric demand causing larger

efferent branches. Nevertheless, in general the sensori-motor demand may be the same

205 F. J. Rühlí - Osteometric Variation of the Human Spine in males and females because it depends on the number of muscle motor units rather than on the size of muscle fibres. One factor to remember, while interpreting osseous dimensions and the possible rçlation to their neural contents, is the fact that the cervical and lumbar enlargements may vary in level even within one species. Therefore, if one finds a different shape of the osseous spinal canal in a certain fossil or skeleton, any interpretation of its altered neural content must be formulated with caution.

To summarize, conclusions on the size and content of the spinal canal, based on the osseous dimensions only, should to be formulated very cautiously. A comparison of

earlier published data of the osseous spinal canal and the measures of this study could be found in Table 14.

F. J. Rühli - Osleometric Variation of the Human Spine 206 c7

Stewart (1962) Shanidar 1 16.0 25.0

Matiegka Predmosti 3 14.0 26.6

Ditto Predmosti 9 12.0 22.0

Ditto Predmosti l4 t4.4 2't.0

Ditto Predmosti 4 13.5 22.5

Earty Medieval Polish (N males=5O, N females= 14.5 23.3 14.2 22.5 Piontek 33/3s)

22.9 Piontek and Zaborowski 12ù-18ù century Polish 14.6 23.8 t5.2

Tomnaga et al Recent Americans (?, N males=4, N females=2, ls.2 26.3 both sexes combined)

Gepstein ef a/. Israeli (both sexos?) 23;7

7) Hasebe Japanese 13.8 23.3 12.9

Europeans t4.7 25.4 14.3 24.3 Aeby

White Americans t4.4 24.8 Lanier

Kø¡dziora et al. Germans (both sexes) 15.9 24.6

22 Stefl

Thomson Recent Europeans (both sexes) t4.7 25

Ditto Recent Bushman l3 2l l3 2I

White Americans 15,5 25.6 t4.4 24.4 Francis

Black Americans 15.5 25.5 14.3 24.4 Ditto

Panjabi et al. Recent Americans (both sexes) t5.2 24.5

Stewart Global samPle (N:20) 12.5 - 17.5 20,0 - 26.0

24.4 Present study (whole samPle) 14.9 25.2 14.3

25.7 Present studY (modern subgrouPs) 15.r 26.1 14.5

Th1

Matiegka Predmosti 3 15.6 24.0

Ditto Predmosti 9 I1.4 2r.4

Ditto Predmosti 14 14.0 23.3

208 F. J. Rühli - Osteometric Variation of the Human Spine

14.9 Dommisse South Africans (both sexes) t3.4

16.3 Thomson Recent EuroPeans (both sexes) 15.7

14 15 Ditto Bushman t4 l5

17.3 15.9 16.6 Present studY (whole samPle) 16.3

t7.7 t6.2 16.9 Present studY (modern subgrouPs) L6.7

Th10

t't.3 Trinkaus Predmosti 3 15.5

16.0 17.4 Matiegka Predmosti 4

14.0 15.0 Ditto Predmosti 10

t5.2 13.6 15.3 Hasebe Japanese 14.3

t6.4 17.5 Aeby Europeans I't.3 18

19 Stefl

t7.2 Lanier White Americans 15.3

15.6 Dommisse South Africans þoth sexes) 13.s

18.2 Pzrjabi et al. Recent Americans @oth sexes) l5,5

t't;7 Thomson Recent EuroPeans (both sexes) 15.3

t6 15 t'7 Ditto Recent Bushman l5

17.3 Marchesi ¿l a/. Swiss (N males= 18, N females=l5, both sexes and 15.8 sides combired)

18.4 15.7 17.3 Present studY (whole sarnPle) 16.2

18.6 16.4 11.9 Present studY (modern subgrouPs) t6.4

L1

22.7 Tri¡kaus DoIni Vestonice 15

25.6 Matiegka Predmosti 3 18.0

22.3 Ditto Predmosti 14 16.0

22.0 Ditto Rom¿no-British (N=?, both sexes) 15.9

2r.3 Porter and Pavitt (1987) Anglo-Saxon (N=?, both sexes) t5.2 t't.l 2r.l Piontek Early Medieval Polish (N males=5O, N r'7.6 22.3 females=45) (1952) 23.4 Huizinga et al. 19* century Netherlands (N:5 I' sex?) 18.0

210 F. J. Rühli - Osteometric Varíation of the Human Spine (1985) Nigerians (N males=79, N females=43) 16.6 15.8 Amonoo-Kuofi

Japanese 16.6 20.3 16 t9.7 Hasebe

(1977) Japanese (N males=59, N females=2l, both sexes t6.2 Kiku'chi et al. combined)

at a/., cited by Japanese (N=?, sex?) t4.3 Takemitsu Kikuchi ¿/ ¿/.

cited by Japanese (N=?) 16.6 t6.4 Nagashima, Kikuchi el ai.

cited by Japanese (N= ?, sex?) t6.4 Tsunematsu, Kikuchi e¡ a/.

by Kikuchi et Japanese (N=?, sex?) r7.0 Hibi, cited al.

cited by Japanese (N:?, sex?) r5.6 Okamoto, Kikuchi e/ a/.

al a/. Israeli (both sexes combined?) 20.8 Gepstei-n

Europeans 18.5 23.9 l8.9 22.8 Aeby

(1995) Koreans (N males=63, N females=27) 15.4 21.5 15.5 20.5 Lee et al.

el ¿/. Swiss (N males= 18, N females=15, both sexes and 17.8 23.5 Marchesi sides combined)

Russians (both sexes) 20 23 Stefl

White Americans 17.2 23.2 Lanier

e¡ ¿/. (1983) Italians (N=63, both sexes) t6.7 2t.7 Postacchini

Indians (N:58, both sexes) 15.0 19. I Ditto

(1987) Àmericans (N=30, both sexes) 17.2 22.1 Berry et al.

Dommisse South Africans (N=25, both sexes) 15.4 20.4

Eisenstein (1977) Caucasoid (N males=78, N females=35) 18.0 23.0 18.0 22.0

Zulu Negroid (N m¿les= 108, N females=54) 16.0 2t.0 17.0 20.0 Ditto

Ditto Sotho Negorid (N males= 106, N females=62) 16.0 2r.0 16.0 20,0

1a ', Àmericans t7.6 17.'l 2t.2 Scoles e, ø/.

al. (1992) Recent Americans (both sexes) 19.0 23.'7 Pzrjabi et

Thomson Recent Europeans (both sexes) t5.7 2t.3

Bushmen (N males=2, N females=l) r5.5 18 t4 20 Ditto

Present study (whole samPle) 17.8 23.7 17.7 22.5

Present study (modern subgrouPs) 18.2 24.4 18.4 23.2

271 F, J. Rühli - Osteometric Variation of the Human Spine L5

23.1 Trinkaus Dolni Vestonice l5

predmosti 3 18.0 27.6 Matiegka .; predmosti 9 15.6 24.3 Ditto

Ditto Predmosti 14 l7 '4 29.9

Ditto Predmosti 4 18.5

26.3 Ditto Predrnosti 10 13.0

Ditto Romano-British þoth sexes) l5'2 25.7

Porter and Pavitt Anglo-Saxon (both sexes) 14'6 25.6

24.9 24.1 Piontek Early Medieval Polish (N males=48, N l'l '3 16.5 females=40/41)

Huizngaet al. 19'century Netherlands (N=51, sox?) 16'9 25.8

30.0 Gepstein el a/. Israeti @oth sexes combined?)

Nigerians 16'0 14.6 Amonoo-Kuofi

Japanese 16'9 26.4 15.6 25 Hasebe

Kikuchi ¿, a/. Japanese (both sexes combined) l5'8

Takemitsu et ¿/. Japanese (sox?) 14'3

Japanese f8 3 16.8 Nagashima

Tsunematsu Japanese (sox?) 16'3

Hibi Japanese (sex?) 18 0

Okamoto Japanese (sex?) 16'3

Marchesi ef c/. Swiss (N males= 18, N females=I5, both sexes and 17.7 21.0 sides combined)

Europeans 19 l 27.5 19.2 25.6 Aeby

Koreans 14'6 25.9 14. t 25.3 Lee et al.

Postacchini ¿/ ¿/. Italians (both sexes) 16'l 24.8

Ditto Indians (both sexes) 14'0 22.8

Stefl

Lanier White Americans 17.4 26.3

16.8 26.0 Scoles ¿t a/. Àmericans 17 '6 25.9

Americans 17.3 26.0 Berry el al. 212 F. J. Rühti - Osteometric Variation of the Human Spine Panjabi. et al. t9.7 27.r Recent Arnericans Donmisse 15.s 24.r South Africans (both sexes) Eisenstein 18.0 26.0 18.0 25.0 Caucasoid

26.0 16.0 24.0 Ditto Zulu Negroid 16.0 Ditto 16.0 25.0 16.0 24.0 Sotho Negroid

11 Thomson 14.3 Recent EuroPeans (both sexes) )7 l4 20 16 Ditto Recent Bushmen Magnuson (1944) Àmericans (?, both sexes?) 12

16.9 26.2 16.9 26.0 Present studY (whole sâmple)

t7.7 26.3 l7.7 26.5 Present studY (modern subgrouPs)

213 F. J. RühIi - Osteometric Variatíon of the Human Spine The intervertebral foramen is an anatomical structure of important clinical value.

a held This will be outlined in depth below as a separate chapter, since it may represent

As already of common scientific interest for anthropologists, anatomists and cliniciancs.

pitfalls. In the highlighted above, the osteometric assessment of this structure has its

to present study, the intervertebral foramen width was measured not only bilaterally,

of the explore any possible side difference, but also on the cranial and caudal surface particular vertebral body. Cinotti et al. (2002) conclude that the measurement of the superior and minimum intervertebral foramen width is a reliable method for the

has been done for the present assessment of the intervertebral foramen dimensions, as it

study. Additionally, Cinotti et al. (2002) state that the impact of the disc space

wet narrowing on the foramen can be shown preferably on dried vertebra rather than

Therefore, by spines, with smaller standard deviations to be found in the first ones'

are exploring the osseous intervertebral dimension, one can assume that the tendencies

similar for cadaver spine diameters and fresh wet spines too. Due to methodologic

difficulties, which have been widely addressed already above, it is difficult to directly

published compare the intervertebral foramen values in the present study with the ones

earlier; see also Table 15.

214 F. J. Ri)hti - Osteometric Variation of the Human Spine

Table 16: Correlation coefficient between individual stature (femur maximum length for present study) and foramen magnum breadth

Sex Röthig (te7t) Present study

Males 0.41 (N:560) 0.0s (N+8)

Females 0.35 6:265¡ 0.18 (N:43)

217 F. J. Rühti - Osteometric Yariation of the Human Spine

30.5 Torguts 36.2

30.3 Malays m 34

28.5 Malays f 32.6

29.9 Australians m 3s.5

29.3 Australians f 34

29.3 Paltacalos m 32.8

28.5 Paltacalos f 35.9

Nakashima (1980:

29:7 MiddleKyushuites m 345

29.8 Kantoites m 35

30.2 North Kyushuites m 36.2

30.3 Yoron-islanders m 35.9

Kikai-islanders m 39.1 30.7

30.2 Shilingol-Mongolians m 37.6

30.3 Fuschen-Chinese m 3s9

29.7 Germans m 3s.3

Present study:

Whole sample m 37.2 32.1

30.0 Whole sample f 35.8

32.4 Modern subgrouPs m 37.3

31.0 Modern subgroups f 35.8

219 F. J. Rühli - Osteometric Variation of the Human Spine The spinous processes, beside their function in limiting extensional forces

(White and Hirsch,l97l), serve as bony lever arms for the back musculature such as multifrdus and spinalis muscles in the lumbar region, whereas the transverse processes act as levers for muscles such as longissimus, psoas major or quadratus lumborum. Both anatomical structures have only rarely been investigated so far with an osteometric perspective, this is in particular true for the spinous process (Schultz, 1961; Cottenll et al., 1986). The transverse process has so far been addressed in limited reports too, all in modern samples only. Furthennore, in the present study the processes often suffered from post mortem damage. This resulted in overall small sample sizes, which led to the exclusion of some of these process measures from the f,rnal data analysis. All this makes it hard to validate the measured dimensions in the present study. A list of earlier published data could be found in Table 18 and 19'

F. J. Rühli - Osteometric Varíation of lhe Human Spíne 220 Table 18: Length of spinous process as a percentage of the sagittal diameter of the vertebral body

Level - Sex Schuttz'(1961) Present study

C3-m 98 % (N:2) 105 Vo (N:37)

c3-f 103 % (N=2) 90 % (N:44)

oÁ C7 -m 214% (N=2) 173 (N:72)

o c7 -f r84% (N=2) 167 (N=65)

221 F. J. Rühli - Osteometric Variation of the Human Spine Table 19: Transverse process width (mm) of various samples

Reference Level / Sample Transverse process width

c3

Hasebe (1913) Japanese - males (N=20) 56

Lanier (1939) White Americans (N=+/-96) 54

Francis (1955) White Americans - males (N=100) 54.9

Ditto White Americans - females (N=27) 50.0

Panjabieta/. (l99la) Recent A¡nericans - both sexes (N =12) 50.3

Ditto Black Americans - males (N=100) s3.3

Ditto Black Americans - females (N=57) 48.9

Cyriax (1920) English (?) - both sexes (N= 70) 53.5

Hasebe Japanese - females (¡=16¡ 50

Present study (whole sample) - males 54.8

Present study (whole sample) - females 50.0

Present study (modern subgroups) - rnales 56.r

Present study (modern subgroups) ' females 51.9

C7

Hasebe Japanese - rnales 7l'5

white Americans '12'5 Lanier

Francis White Àmericans - males 72'4

Ditto White Àrnericans - females 65'4

Parjabi et al. Recent Americans (both sexes) 66'6

Ditto Black Àmericans - males 70'2

Ditto Black Americans - females 64'5

Cyriax English - both sexes 68'0

Hasebe Japanese - females 68'9

Present study (whole sample)'males 66'2

Present study (whole sample) - females 54'6 222 F. J. Rühli - Osteometric Variation of the Human Spine Present study (modern subgroups) - males 66.2

Present study (rnodern subgroups) - females 52.9

Thr-;

Japanese - males 74.8 Hasebe

White Arnericans 78 Lanier

English (?) - both sexes '74;7 Cyriax

Recent Americans - both sexes 7< 1 Patja,bi et al. (l99lb)

Japanese - females 6s.5 Hasebe

Present study (whole sample) - males 78.0

Present study (whole sample) - females 70.9

Present study (modern subgroups) - males 79.1

Present study (rnodern subgroups) - females 72.5

Th6

Japanese - rnales 62 Hasebe

Cotterill et a/. (1986) Canadians - both sexes? (N =10) 55.7

White Americans 6t Lanier

Recent Americans (both sexes) 61.3 Panjabi et al.

English (?) 63.6 Cyriax

Japanese - females 54. I Hasebe

Present study (whole sample) - males 65.1

Present study (whole sample) - females 59.7

Present study (modern subgroups) - mâles 65.5

Present study (modern subgroups) - females 60.9

Th10

Japanese - rnales 56.3 Hasebe

\ühite A,mericans 60.8 Lanier

Recent Americans (both sexes) 58.4 Parjabi et al.

English (?) 5 8.5 Cyriax

223 F. J. Rühti- Osteometric Varíation of the Human Spine Japânese - females 49.4 Hasebe

Present study (whole sample) - males 60.7

Present study (whole sample) - females 55.0

Present study (modern subgroups) - males 63.0 - Present study (modern subgroups) - feniales 58.1

L1

Japanese - males 67 Hasebe

lYhite Arnericans "13.1 Lanier

(1992) Reeent Americans (both sexes) 71.2 PalLjabi et al.

English (?) 72.6 Cyria*

Japanese - females 61 Hasebe

Present study (whole sample)' males 73.0

Present study (whole sample) 'females 64.6

Present study (modern subgroups) ' males 75.1

Present study (modern subgroups) - females 68.3

L5

Japanese - males 88.8 Hasebe

\Vhite Americans 92.6 Lanier

Recent Americans (both rexes) 92.5 Parjabi et al.

English (?) 86.0 Cyriax

Japanese - females 82.4 Hasebe

Present study (whole sample) - males 85.2

Present study (whole sample) - females 78.0

Present study (modern subgroups) - males 91.s

Present study (modern subgroups) - females 84.5

224 F. J, Rühli - Osteometric l/ariøtion of the Human Spine Anotherfactor-toinvestigatewhiledoingmorphometricresearchisapossible

couid be found for the intra-individual side difference. For example, no side difference with most earlier reports pedicle dimensions in the present study, which is consistent Kothe et al., 1996). As (Marchesi et al.,lggg; Bantà et al., 1989; Xu et al., 1995;

did not show any side difference another exemplary measule, the vertebral body height (1883)' who linked the higher in the present study, unlike in the report by Anderson

bigger weight of the internal organs values of the right side of the vertebral body to the

on this side.

Tosummarize,ifonecompafestheosteometricdataofthepresentstudyofboth' with earlier published the whole sample as well as the selected modern samples, of the present study fall within the measures, it can be seen that most of the measures e'g'' the transverse process wide range of spinal dimensions. Some exceptions are published earlier' on the widths at c7 ,which in this study are smaller than the measures are in this study larger than the other hand, the pedicle dimensions at Th6 and Th10 of the human spine, as e'g'' seen in ones published so far. Since the overall variability quite large, these outliers of the the above shown dimensions of a global sample, is

of this variation or simply be caused present study may de factojust represent extremes

by minor methodologic differences'

Variation of spinal morphometry due to sex ønd age

Both,sexandindividualugearekeyfactorscontributingtothevariabilityin

above, and, therefore, were major osteometric measurements, as already highlighted

were elaborated specifically on the issues addressed in the present study' Both factors

and age' two modern samples with historically known individual sex

225 F. J. Rühti - Osteometric Variation of the Human Spine In the study presented here, males show for most vedebral measurements

significantly higher values such as vertebral body height ot transverse process width. In the most similar microevolutionary study of the human vertebral column, Jankauskas

(1994) estimated the overali influence of sex to be about 30% - 40Yo of the variability of

vertebral column, with age having an impact of just about 5% - 8%' Sex was a

significant factor especially for transverse diameters. He reports for the cervical,

thoracic and lumbar spine a signihcant sexual dimorphism for the vast majority of

osteometric measurements, consistent with the findings of the present study. Sex was

also not a major contributor towards the occurrence of spinal pathologies in the

archaeologic sample examined by Jankauskas (1992). Nevertheless, males have e.g., a

significantly longer thoraco-lumbar spine (Gozdziewski et al.,1976). These factors, due

to restriction on selected vertebral levels and non-pathologic spines, could not have been

explored in the study presented here. Despite the fact that Huizinga et al. (1952),

surprisingly, do not consider possible age and sex estimations as further factors

influencing their hndings on lumbar spinal canal dimensions in historic skeletons, most

authors agtee that sex seems to influence the spinal morphometry (Piontek, 1973;

Larsen and Smith, 1980a; Larsen and smith, 1980b; Hermann et a1.,1993). The results

of the study presented here also support this view'

With respect to neural pathways of the spine, the results of the present study are

notable. No signifrcant sexual dimorphism can be found, basically, for the osseous

outline of the neural pathways, unlike in the other spinal osteometric dimension such as

e.g., the vertebral body height. This notable absence of larger neural dimensions in

males has already been reported in similar way earlier (Eisenstein,1977; Porter et ø1.,

226 F. J. Rühli - Osteometric Variation of the Human Spine 1978b). For example, Porter et al. (1980) found, in an ultrasound based study, larger neural canals in females than males, especially in the subgroup of young adults. They mention a possible higher amount of epidural fat as one possible cause and the likely advantage in case of pregnàncy-related mechanical stress to be responsible for this size difference. The lack of sexual dimorphism in neural canal dimensions does not mean there is no sexual difference in its shape, as earlier shown for a different prevalence of a trefoil shaped lumbar spinal canal (Eisenstein, 1980). Females show larger osseous and non-osseous spinal canal cross-section areas, but smaller neural tissue cross-section areas (Hasue et al., 1983; Kikuchi et al., 1984). This may make males, especially atL5 where the difference is most obvious, more vulnerable to any pathologic conditions

(Hasue et al., 1933). Surprisingly, there is even one dimension in the present study, which is significantly bigger in females than in males, the right caudal intervertebral foramen width on level L5. This might have clinical implications as will be discussed in depth further below. Similar hndings were also reported for the intervertebral foramen and spinal nerve root dimensions (Hasue et al., 1983; Kikuchi et al., 1984)'

Additionally, Hermann et at. (1993) could not find a difference of the subarachnoid space in relation to sex. With regard to nerve root size, one has to be aware that females have in absolute terms the same measures, which makes them relatively to body weight even bigger than in males, as shown by Dunn (1912) for rats at least.

The particular impact of aging has been explored in the study presented here as well. Based on the modern samples with historically recorded individual age, both sexes

show alterations of spinal and long bone morphology with aging. In the study presented

F. J. Rühli - Osteometric Variation of the Human Spine 227 males. This may be here, aging was found to contribute significantly de facto only in due to their larger sample size, especially in the modern sample'

aging; In the present study, the neural pathways do not change significantly with

dimensions' this is unlike earlier reports on age-related alterations of neural osteometric

cord diameter to Humphreys et at. (1998) describe an increase of the ratio of spinal

They also detected spinal canal diameter in the early adult asymptomatic cervical spine.

20 30 years and a an increase of the C6 I Cj foraminal width in the age group of to

the age of 50 years slight decrease followed by another increase later for the older below

(Humphreys et al',1998)' category, unlike the steady decrease in symptomatic patients

The osteometric dimensions unfortunately can give just a glimpse of the age-

leads a decreased related alterations of their neural content. For example, aging to

the spinal nerve number of myelinated fibres and an increase in connective tissue in

to be no change roots (Dunn, l9l2; Corbin and Gardnet, 1937)' However, there seems but in in the dorsal root / ventral root ratio with age (Corbin and Gardner, 1937), in the growing rats the increase in nerve fibres was for a longer time and more intense

dorsal nerve root (Dunn, I9l2).

Furthermore, one has to be aware that the stable dimensions of the neural

structures pathways in the present sfidy de facto Íwresent a relative decrease of these

pedicle height' with age, since other neighbouring osseous structures, such as the significance as well is apparently increase with age. whether this may have clinical capacþ, which is doubtful. weisz and Lee (1983) found that the spinal canal reserve

and of the spinal cord, the difference between the sagittal diameters of the osseous canal

to decreases of decreases with age, making the elderly apparently even more wlnerable

228 F. J. Rühti - Osteometric Variation of the Human Spine size. Nevertheless, it is still debated if lumbar neural canal in the spinal canal ".g., general were becoming bigger (Clark et a\.,1985) or smaller (Porter et a1.,1980;Lee et al., 1995; Tatarek, 2001) with aging and some reports could not f,rnd a clear link between spinal cord alteratior, *d individual age (Elliott,1945; Bailey, 1953; Legg

and Gibbs, 1984). In a study by Lee et al. (1995) a significant influence of age on spinal morphometry, as shown for a decrease in lumbar mid-sagittal and transverse spinal

canal diameters, occurred only after the age of 60 years. Therefore, this factor due to the

aveÍage low mean age in archaeologic samples may not be that relevant. The average

age in the historically recorded modern samples is in the present study even below 50

years.

In the present study, the found age-related trends of spinal morphometry,

signifrcant after Bonferroni's correction only in males, are some pedicle heights and

sagittal and transverse vertebral body diameters. The last one is consistent with earlier

reports (Jankauskas, 1992; Jankauskas, 1994), explained as a possible effect of

degenerative changes (Jankauskas, 1994), but a decrease of anterior vertebral body

height with age (Jankauskas, 1992) could not be found in the present study.

The increase of the vertebral body and pedicle diameters with individual age in

the present study seems to be due to a general increase in robusticity in the elderly, a

remodelling resulting in a surplus deposition of bone, which most likely does not affect

the osseous outline of the neural pathways. The robusticity generally changes with age,

most prominent for the measurements of the long bone shafts, as reported in the

literature (Pfeiffer, 1980) and possibly as a physiological reaction to compensate for loss

of stiffness due to a general decrease in bone mass, especially in women (Pfeiffer,

F. J. Rühli - Osteometric Variation of the Human Spine 229 the modern age-recorded 1930). In the present study, the long bones, as tested for

expressing an increase in samples, showed for males such an increase in robusticity,by

females, only femur length circumference but no signif,rcant change in length, but for

decreased signifi cantlY

Inter-p opulational variations of spinal morphometry

highlighted The inter-populational variations of spinal morphometry, as already

McCotter, 1916; by various authors (wetzel, 1910; Hasebe, i913; Thomson, 1913;

Rasmussen, 1938; willis, 1923; Stefko, 1926;Martin, 1928; Stewart,1932; Lassek and 1955; Bomstein and Matiegka, 1938; Wood-Jones, 1938; Lanier, 1939; Francis, 1976; Eisenstein, 1977: Peterson, 1966; Piontek and Budzynska, 1972; Ericksen, Amonoo-Kuofi, 1985; Eisenstein, 1980; Tibbetts, 1981; Postacchini et al', 1983;

et al.,1995; Tatarek, Nakashima, 1986; Ross e/ al.,l99l; Jason and Taylor,1995;Lee

samples belong to a 2001) will hardly apply in the present study, since all selected

modern the European samples Centrai-'Western European g1oup. Nevertheless, the more

variability, at least as are, the more likely decreased the inter-group morphological

Whether this is the case for the shown for cranial characters (Henneberg et a1.,1978)'

further investigated' spinal morphometry as well would also be worth to be

Relation of geography and society to spinal morphometry

human body' Various environmental factors influence the morphometry of the

selected morphological For exampie, the geographic latitude alters the expression of lateral internal thoracic traits in humans, such as biiliac breadth (Ruff, 1994) or the

230 F. J. Rühli - Osteometric Variation of the Human Spine artery (Surtees et al.,1989a; Surtees et al., 1989b; Henneberg, 1992)' In the present study, all individuals come from similar geographic latitudes, approximately 45'N - 49o

N. This should ruie out major influences of latitude on the spinal morphometry.

Furthermore, the unique situaiion of the more alpine populations in Switzerland, as for example of the Chur sample, has already been highlighted above'

From a cultural point of view, the samples presented here reflect various historic transition periods, from prehistoric hunting and gathering populations, such as Upper

Paleolithic, to a more sedentary agricultural dispersed life-style, such as Neolithic and

Brotue Age, semi-urban and urban societies in Medieval times and, finally, post-

industrialization communities.

The influence of changes in European life style and its effect on human gtowth,

morphological characteristics, morbidity and mortality has been studied in numerous

reports (Henneberg et al., 1978; Lewis,2002).In general, two major morphologically

distinguishable groups are known for the European Holocene; a Southern-Western

European population type and Northern-Eastem series (Schwidetzky, 1967;

Schwidetzþ,1972; Schwidetzky and Rösing, 1976; Rösing and Schwidetzky, 1977;

Rösing and Schwidetzky, 1981; Schwidetzky and Rösing, 1984: Schwidetzky and

Rösing, 1989). The geographical distribution and the inter-populational difference

decreased during most time periods (Schwidetzky,1967; Schwidetzky,1972). For some

dates regional differences became more apparent towards more modern times and

Rösing and Schwidetzky (1981) name increased social differentiation in the form of

religious or urban versus rural locations as possible factors. Furthermore, a remarkable

closer similarity of the population subtypes within the Westem samples than for the

F. J. Rühli - Osteometric Variation of the Human Spine 237 Eastern series has been described (Rösing and Schwidetzky, i977; Schwidetzþ and

Rösing, 1989). This is of interest since the selected series of this work, withthe single exception of the Hainburg material, would most likely belong to the Western population clusters and small inter-populuìiorrut differences help to analyse the various groups.

The vast majority of the selected individuals in this work originate from inland non-coastal ecozones. The only exception would be the French Mesolithic individuals'

Changes in Upper Paleolithic to Mesolithic in Europe have been of socio-cultural and

ecological nature, with an increased population density, with a decrease in nomadic

lifestyle, an increased resource reliability and food abundance but also an increased

technological sophistication, all factors contributing to an ecological framework relying

on the interaction between resource-stress and humans (Hayden, 1981)'

Body and especially limb morphology seem to be influenced by various factors

such as gene flow, transmitted by a population movement from Sub-Saharan Africa

towards Europe - and which resulted in altered metabolic demands and vasomotoric

adaptation to a cold environment - or, finally, stress due to physical activity (Trinkaus,

19gl; Holliday,1996; Holliday, 1997; Holliday,1999). The importance of these factors

for the spinal column morphology in particular cannot conclusively be said at this stage.

At least it is well known, that limb proportions changed in Europe from a more sub-

shape Saharan Afücan type in Early Upper Paleolithic to a more modem European body

in the Late Upper Paleolithic (Trinkaus, 1981; Holliday, 1996; Holliday, 1997;

Holliday, 1999). Mathers and Henneberg (1996) suggested a changing of relative trunk

size and lower limb proportions to be represented in the found different trends for

hominid body height and weight within the last 4 millions of years. Since they found no

such divergence of trends in Homo sapiens body weight and height for the last 32'000

232 F. J. Rühli - Osteometric Variation of the Human Spine in this time frame, years, they propose poisible different microevolutionary trend acting which would be of particular interest for the present study; such microevolutionary trends will be elaborated separately further below'

must be The impact of physicaily demanding life-style on the vertebral column

historic transition taken into account as well. The decrease in robusticity during such life-style periods was most likely related to adaptations to physically less demanding by (Larsen, 1980; Larsen, 1981; Larsen, lgsz). Nevertheless, one interpretation

patterns, possibly schwidetzþ (1962) arguing that specific character and behavioural

linked with level of gracilisation, could have been seiective in such changing

environment, seems to stretch the case' physical activities Another physical factor, the age of commencement of adult

factor ofbone robusticity has so far been supported to a variable extent as an etiological agricultural societies show alterations (Bridges, 1993; Knüsel, 1993). Apparently, some

degenerative bone disease than their a higher bone robusticity but a lower prevalence of

heavy labour in the first life- hunter-gatherer counterparl as a result of juvenile onset of

young members in a style group (Knüsel, lgg3). The early physical involvement of

skeletal robusticity and settled society would allow these individuals to have a higher

to degenerative osseous plasticity later in life and, therefore, less likeiy to be vulnerable for the morphometry of alterations (Knüsel, 1993). This seems in particular reasonable

the vertebral column.

Not only the selection of a clinical or historic spinal sample, but also its

alterations in spinal geographical, environmental and ethnic background contributes to

are always applicable morphology; therefore, normativ e data for spinal morphometrics

only (Ross et al',1991; Hermam' et to a certain degree for a confined geographical area

233 F. J. Rühti - Osteometric Variation of the Human Spine present study' Further bio-socio- al., 1993). This is most likely also true for the

reveal a deeper insight into their archaeological studies on the examined samples would its particular impact on the particular cultural situation, which are crucial to better assess

spinal osteometric values.

Influence of stature and body size on spinal morphometry of the spinal column' its If one investigates microevolutionary and secular trend

as well' It is well known that the individual dependence on stature needs to be assed individual stature (Dwight' particular spinal morphology and length is a function of

et al',1976; Galiagher et al'' 1894; Hasebe, 1913;Fully and Pineau, 1960; Gozdziewski

1988;Minneetal.,1988).Acorrelationbetweenindividualvertebralbodyheightor

(Fully and Pineau' 1960; Tibbetts' pedicle height and stature has been described earlier findings can be reported for 1981; Galla gher et al., 1988; Scoles et a1.,1988). Similar

the study presented here. main diameters in ln the present study, most of the vertebral body height and

see also appendix 9' In males' both sexes correlated with individual femur length; heights show such a significant selected transverse process widths and pedicle foramen dimensions and correlation as well, whereas in females, selected intervertebral

pedicle heights do.

some of the earlier studies To assess individual stature from spinal dimensions

equation consisting of lower limb propose for accurate individual stature estimation an

spine such as the lumbar region in long bones such as femur or tibia and parts of the and Pineau, 1960; Tibbetts' 1981)' case of just partial skeletal preservation (Fully

234 F. J. Rühti - Osteometríc Variation of the Human Spine

in Europe (Frayer, 1980; Frayer, 1981; Frayer, 1984; Jacobs, 1985b). Alterations, such as the reduction in masticatory and gastrointestinal tract as well as in the musculoskeletal system interfered with the supposed body shape changes (Henneberg,

200 I a).

The natural selection of body size is influenced by long-term genetics, such as constant mutations, genetic interbreeding or gene pool drifts, and in short-term more by direct environmental factors. Frayer (1984) postulates smaller bodies being energetically more economical and, therefore, been naturally selected in times of lack of ressources.

Gracilisation seems to be created by technological improvement during human evolution. Smaller bodies are more fit in terms of food efhciency under conditions of decreased demands for physical strength and robusticity (Frayer, 1981; Henneberg and

Steyn, 1995).

Human skeletal morphology reflects its genetic and environemental influences,

as already discussed above. The skeletal robusticity alterations seem to be rather

dependent on long-term and repetitive mechanical forces, whereas degenerative changes

more likely seem to be related to traumatic or intense but rare impacts (Bridges, 1991).

The question, therefore, remains at least partially unsolved how far changes in life-style,

as seen from a hunter-gatherer society towards an agricultural community, influence

bone morphology or what other factors contribute as well.

Human postcranial robusticity is undergoing various changes (Ruff et al.,1993;

Trinkaus, IggT). Alterations in biomechanical loading, hormonal or genetic adaptations

control bone remodelling especially of the diaphyseal bone (Ruff et a1.,1993; Trinkaus

et al., 1994; Tntkaus, i997). The humerus is an excellent long bone to show any

F. J. Rùhli - Osteometric Variation of the Human Spine 237 plasticþ, because the humerus does not show any impact from locomotion and systematic influences such as nutrition will appear symmetrically (Trinkaus et al.,1994;

Trinkaus, lggl).In the present study, the humeral changes are found to be consistent, in

males and females, with the femoral ones.

Gracilisation occurred as described above in Europe, Australia and East Asia

(Brown, Igg2), whereas in Africa and certain regions of Australia its extent is still

pattern debated (Henneberg and Steyn, 1993; Pretty et a\.,1998). The extent and precise

of the European gracilisation is sti1l debated, as widely outlined above. The general

trend in decrease of robusticity, apparent from the Late Paleolithic to more modern

times (Formicola and Giannecchini, lggg), is at least in some of the selected individuals

originating from Central Europe not observable. Whereas the gracile Téviec-island type

individuals, among others all of the selected Hoëdic and Téviec samples, in general

follow the trend of postcranial gracilisation and decrease in individual stature, the more

robust Téviec-continental t¡pes, among others the Gramat individual in the samples of

the present study, do this to a lesser extent (Vallois and de Félice, 1977)-

In general, the selected samples for the present study may not be representative

enough to highlight in particular the microevolutionary alterations of long bone

morphology, since this was not the main issue of this work. Tables 2l -24list earlier

reported humerus lengths, femur lengths femur head breadth, femur mid-shaft

circumference and bi-iliac width, whereas Table 25 lists estimated statures of various

historic European samples. A summary of these values could be found in Figures 22 -

25. Figure 26 shows the means of the measured long bones in the present study.

238 F. J. Rühti - Osteometric Variation of the Human Spine

Table 2l : Femur length (mm) and femur head breadth (mm) of various samples

Sample Femur Femur tr'emur Reference length - length - head males females breadth

Ruff(1994) Early Upper Paleolithic, modern -É[. 46t 48.1 søpiens (N:i 1, both sexes)

La Chapelle-aux-Saints (m) 433 Ditto

Predmosti (m) 455 Ditto

European Neandertals (N:5, both sexes) 434 51.'7 Ditto

Late Upper Paleolithic, modern .É[. 434 46.7 Ditto søpìens (N:4, both sexes)

European Upper Paleolithic (N 471 422 Frayer males:l7, N females=5) (1e81)

European Mesolithic (N males:2O, N 444 409 Jacobs females= 17) (1e8sb)

European Mesolithic (N males:16, N 43s 404 Frayer (1e8 females:13) 1)

Pre-agricultural Americans (1000 BC- 449 434 45.5 (m) / Larsen (1e81) 1150 AD) (N males:9, 14; N females:l9, 41.1 (Ð 31)

Agricultural Americans (after 1150 AD) 448 416 a3.8 (m) / Ditto (N males:47, 58; N females=54,61) 3e.0 (Ð

American Whites (N males:255, N 473 430 Trotter and females:63) Gleser (res2)

244 F, J. Rühti - Osteometric Variation of the Human Spine European samples Table22: Femur mid-shaft circumference (mm) of various (Jacobs, 1985b)

Femur circumference Sample

93 Upper Paleolithic - rnales (N=16) 78 Upper Paleolithic - females (N=8) 94 Mesolithic - males (N:16) 80 Mesolithic - females (N:15)

241. F. J. Rühli - Osteometric Variation of the Human Spine

Table24:. Estimated statures (cm) of various European samples

Sample Stature Reference

Neandertals (N males=4) 163 Various studies, cited by Martin (1928)

La Chapelle-aux-Saints (N malrl) t64 Ruff(1994)

Mean male archaic Homo sapíens t67 Ditto

Mean male early modern,ÉIomo sapíens t77 Ditto

Upper Paleolithic (N males:20) t74 Frayer (1984)

Upper Paleolithic (N females:9) 1s9 Ditto

Italian Upper Paleolithic (N males: 12) t64-l'18 Formicola (1983)

Italian Upper Paleolithic (N females:3) 153-168 Ditto

Early Upper Paleolithic (N males=10) t74 Frayer (198 1)

Early Upper Paleolithic (N females=5) 161 Ditto

Late Upper Paleolithic (N males=10) t74 Ditto

Late Upper Paleolithic (N females:4) t57 Ditto

Late Paleolithic - Veyrier (N malrl) t69 Pittard and Sauter

Mesolithic (N males:26) 165 Ditto

Mesolithic (N females:l5) 154 Ditto

Italian Mesolithic (N males:10) t62-172 Formicola

Italian Mesolithic (N females=4) 150-l5l Ditto

Late Upper Paleolithic (Central Europe, N males=7) 166 Formicola and Giannecchini (1999)

Late Upper Paleolithic (Central Europe, N females=7) 155 Ditto

Mesolithic - Téviec (N males:7) t59-162 Pittard and Sauter (1945)

Mesolithic - Téviec / Hoëdic (N males=I0) 161 Formicola and Giannecchini

Mesolithic - Téviec / Hoëdic (N females=12) 151 Ditto

Mesolithic - Gramat (N male=l?) 165- 166 Ditto

Mesolithic - Birsmatten (N male=l) 160 Sedlmeier and Kaufrnann (1996)

Mesolithic (Western Europe, N males=96) r63 Formicola and Giannecchini

F. J. Rühli - Osteometric Variatíon of the Human Spine 243 Mesolithic (Western Europe, N females=72) 151 Ditto

Mesolithic (N males:4 l) 168 Frayer (1984)

Mesolithic (N females:26) 156 Ditto

Mesolithic (N females= 5) 153 Formicola and Franceschi

Neolithic (N males:62) 166 Frayer (1984)

Neolithic (N females:46) t54 Ditto

Neolithic - Italy (N males=24) 162 Formicola

Neolithic - Italy (N females=17) 151 Ditto

Neolithic - France and Belgium (N males=127) t63 Pittard and Sauter

Neolithic - France and Betgium (N females:53) l5l Ditto

Eneolithic / Bronze Age -Italian (N males:14) 164 Formicola

Eneolithic / Bronze Age -Italian (N females:14) 153 Ditto

(2002) Pompeiians - 79 A.D. (N males:127) t63-169 Henneberg and Henneberg

Pompeiians - 79 Ä.D. (N females:I45) t52-156 Ditto

Bajuwars - 400-800 A.D' (both sexes) t7 t-173 Various studies, cited by Wurm (1982)

Francs - 500-800 A'D' (N males=47) 166 Pittard and Sauter

Francs - 500-800 A'D' (N females=l6) 152 Ditto

Francs - 400-900 A.D. (both sexes) t7 r-173 Various studies, cited by Wurm

Alemanns - 400-800 A'D. (both sexes) 170-l'74 Various studies, cited by Wurm

Alemanns - 400-800 A.D. (Swiss, both sexes) 172 Ditto

Àlemanns - Swiss (N males:750) t69 Pittard and Sauter

Alemanns - Swiss (N females:455) 158 Ditto Ditto Alemanns - Swiss, 700 -1200 A.D. þoth sexes) 165-170

French - 900-1100 A.D. (N males:140) 166 Pittard and Sauter

French - 900-1100 A.D. (N females=46) 156 Ditto

French, Medieval Ages (N males=294) 166 Ditto

French, Medieval Ages (N females=101) 156 Ditto

Medieval (N males=41) 169 Frayer (1984)

Medieval (N females=46) 156 Ditto

244 F. J. Rühtí - Osteometric Variation of the Human Spine Southern Germans, 1180-1400 A.D' 166-168 Various authors, citedby Wurm

Rural Polish, 1200-1400 A.D. (males) 172 Henneberg et al, (1984b)

Rural Polish, 1200-1400 A.D. (females) 159 Ditto

Rural Polish, 1400-1600 A.D. (males) t70 Ditto

Rural Polish, 1400-1600 A.D. (females) 161 Ditto

Rural Polish, 1600-1700 A.D. (males) t7t Ditto

Rural Polish, 1600-1700 A,D. (females) 160 Ditto

Rural Polish, 1700-1900 A.D. (males) t7l Ditto

Swiss conscripts, 1500-1650 Ä.D. t64-168 Various authors, cited by Wurm

Lithuanians, 1o Millennium A.D.(N males:24) r74 Jankauskas (1994)

Lithuanians, 1o Millennium Ä.D. (N females:l6) 161 Ditto

Rural Lithuanians, 2"o Millennium A.D. (N males= 168 Ditto 62)

Rural Lithuanians, 2"0 Millennium A.D. 157 Ditto (N females:29)

Urban Lithuanians, 2"0 Millennium A,D. r67 Ditto (N males:205)

Urban Lithuanians, 2"0 Millennium A.D. r56 Ditto (N females:180)

Modern (fernales) t70 Frayer (1984), with data from Eveleth and Tanner ( I 976)

Modern (males) 174 Ditto

Modern South Àfricans of European extraction 179 Henneberg and van den Berg (1990) (males)

Modern South Africans of European extraction 165 Ditto (females)

F. J. Rühli - Osteometric Variation of the Human Spine 245 Þrl \ mm mm s- tc o) co (¡ n f\) f\) Iu tu eeßß8EBBõc¡õino(¡o<¡r (D 883383Ë N) I N) Upper Paleolithic (Jacobs) o Upper Paleolithic (Jacobs) (\ o O'>o> *_ o tss ELJ Upper Paleolithic (FraYer) -\ CD tsJ. Upper Paleolithic (FraYer) (DH o oÕFiÉ .o æE Mesolithic (Jacobs) G x(Dõa Mesolithic (Jacobs) c Ê (DdË iB= 3 a/lè o o-oo5 a^ Ø E \- Fi P+) U, a:_ < õ Mesolithic (FraYer) G 9Þ Mesolithic (FraYer) o CD{. G¡ l.)y (Í¡ 9CA ) o Americans u). Pre-agricultural Americans il ¿ Pre-agricultural il 3 o (Larsen) o d (Larsen) o Ê) Þ. b o b o c) \t ø v, Þ x \¡ o. + x Americans o Agricultural Americans + H Agricultural æ, b) (Larsen) nq oo' (Larsen) ilg (D 9F r9 (¡)ON o! É ¡ b, T. a) ßp l\)-T Þ (oÀ iÞ ul it Euro-Americans (Trinkaus) (o Euro-Americans (Trinkaus) o\ l\) -.J x o) o + X (¡) + -u, (Þ fÞ O è Flo' (¡ o) White Americans (Trotter and White Americans (Trotter (¡Þ Gleser) and Gleser) Èl.J o, ! Þfl :- oc mm mm Étsl (D ÀÀ.LÀèÞÞÞè Þ N) EFäd8U8Eå ÈàåâÈâââ5$å I !P European Upper Paleolithic (Jacobs) oÞ1 ? Upper Paleolithic (Jacobs) =(D (D sæ Þ) ã' oi + s-o (D' :.o U Ècø É Upper Paleolithic (Frayer) Þ1 (Frayer) European :(D=.(D Upper Paleolithic .o Fl (Þ Þ) iJ õÞa 0q .Tl .Tt '*(D o SNr o o SH H) 3 3 (Jacobs) c European Mesolith¡c (Jacobs) c Þ) Mesolithic ,a- il. \ o ó- o ='G É ct) (o (o

U) European Mesolithic (FraYeQ il ts1 Mesolithic (FraYer) o o 3 .o 3 b A¡ Þ ß¡ è i'J o Þ- o x{ U, H iJ + o o -9 Ê. Pre-agricultural Americans (Larsen) b Pre-agricultural Americans (Larsen) CD N) r8 rt å o@ x(,) boä U) (o f\) Þ) + ó! it J) O io rd -t N) il (o N) CD o) (t)o o N) x Agricultural Americans (Larsen) iu x Agricultural Americans (Larsen) + (o s+ o' @ çn @ ã (o o o { ¿n -{(¡ 6) o x Ep + Þ Modern White American (Trotter and (D Americans (Trotter and Gleser) (¡l\) (D \ Gleser) NJ\È @ 180 y = -0.0002xs + 0.æ58f - 1.0748x + 173.53 R2 = 0.3834

175

170

E (J

tbð

160

155

Figtre 24: Estimated male statures of various historic and modem samples, for complete references see Table 24.

l) Early Upper Paleolithic 20) Neolithic

2) Mean male a¡chaic /1omo sapiens 2l) Neolithic - Italy

3) Mean male early modem Homo sapiens 22) Neolithic - France and Belgium

4) Neandertals 23) Eneolithic / Bronze Age -Italian

5) La Chapelle-aux-Saints 24) Pompeiians - ?9 A.D.

6) Upper Paleolithic 25) Francs - 500-800 A.D.

7) Italian Upper Paleolithic 26) Alemanns - Swiss

8) Late Upper Paleolithic 27) Lithuanians, l" Millennium A.D.

9) Late Upper Paleolithic 28) French - 900-1100 A.D.

10) Late Upper Paleolithic 29) French - Medieval Ages

I l) Mesolithic 30) Medieval

l2) Italian Mesolithic 3l) Rural Polish 1200-1400 A.D.

l3) Téviec 32) Rural Polish 1400-1600 A.D.

14) Téviec / Hoëdic 33) Rural Polish 1600-1700 A.D.

15) Gramat 34) Rurai Polish I700-1900 A.D.

16) Birsmatten 35) Rural Lithuanians, 2*Millennium A.D.

17) Yeyrier 36) Urban Lithuanians, 2"0 Millennium A.D.

18) Mesolithic 37) Modem

19) Mesolithic 38) Modem South Africans of European exhaction

F. J. Rühli - Osteometric Variation of the Human Spine 248

J - ! c c o 6 o o > 300 > 280 400 500 500 7

BPGROUP BPGROUP

A B

460

J L I= c c d 6 o o > 430 400 3 00 4 00 5.00 2.00 300

BPGROUP BPGROUP

C D

Figure 26: Means of measured long bones by time periods in the present study (l:Paleolithic, 2:Mesolithic, 3:Neolithic, :Bltorø;e Age, 5:Early Medieval, 6:Late Medieval, 7:Modem)

A) Humerus length (males), B) Humerus length (females) C) Femur length (males), D) Femur length (females)

F. J. Rühli - Osteometric Variation of the Human Spine 250 demarcation between Western and Eastern European sampies (Formicola and

Giannecchini, 1999). From a cultural point of view the Upper Paleolithic should not be regarded as a simple uniform pan-European period (Straus, 1995). ln terms of skeletal records, this may be different. However, a conclusion, based on the samples included in this study, cannot be reached. The limitation in the current sample to Central and

Western European origin avoids some possible problems, though the findings will be

only applicable to the Westem European region. Nevertheless, there is an obvious need

of further studies to focus on a possible inter-regional difference of the spinal

morphology.No major intra-regional differences in stature within the Westem European

Late Upper Paleolithic and Mesolithic samples have been reported, with in general

lower stature values for the'Western group than for their Eastern European counterparts

(Formicola and Giannecchini, 1999). The large stature of the Early Upper Paleolithic

Europeans could be explained by various factors (Formicola and Giannecchini, 1999).

Funerary behaviour could indicate a bias towards socially higher and possibly male

individuals (Frayer, 1981), but this seems to be rather unlikely.

Climatic adaptations reflected in the found high values of the Early Upper

Paleolithic individuals, as also found in the present study, are even more controversial.

According to the ecologic-adaptive rules by Bergmann (1847) and Allen (1817), stating

that individuals living in cold climate have on average shorter limbs in relation to their

trunk and have larger body mass, the Ice Age maximum would favour more bulky

individuals. For example, the Neandertals seem to be on the average 10 cm shorter, but

3.5 kg heavier than their early anatomically modern human counterparts (Ruff, 1994),

which has been interpreted as a left-over of ancient climatic condition (Formicola and

Giannecchini, 1999). Some interpret the alterations of European body shape and limb

F. J. Rühti - Osteometric Variation of the Human Spine 252 proportions, in particular since the Early Upper Paleolithic, as being related to environmental and genetical influences (Trinkaus, 1981; Holliday, 1997; Holliday,

1999), while others argue that the importance of the climate for the morphologic alterations described for the European Paleolithic-Mesolithic transitions period to be of lesser significance (Frayer, 1981). Nevertheless, in this study all samples come from a temperate Central European climate and from similar latitude, as already outlined

above, so climatic changes would have been most likely similar for the various samples

and of known Central-Western European type.

Evolution of hunting technique, such as the use of spears and, later most likely in

Mesolithic times, of the bow, together with the disappearance of the megafauna, could

have had an impact of body morphology, such as skeletal robusticity and individual

stature from the Late Pleistocene onwards. The increased hunter-game killing distance

by using more developed techniques lowers apparently the human need for high

robusticity and long upper limbs (Frayer, 1981). Furthermore, the particular importance

of the prey size on the development of sexual dimorphism in terms of individual stature

has been mentioned as well (Frayer, 1980).

Changes in nutrition, such as decreased protein intake due to increased

population density, and natural selection favouring longer limbs couid be additional

interfering factors (Wurm, 1982; Formicola and Giannecchini, 1999). The possible

particular nutritional situation of the selected samples in the present study, mostly from

Southern Germany and Switzerland has already been addressed above, based on the

important study on the impact of the protein intake on human morphology (Wurm,

1982). Nutritional influences were also controversially discussed as possible etiologies

of human morphology by various authors (Frayer, 1981; Larsen, 1981; Trinkaus, 1981).

F. J. Rühti - Osteometric Yariation of the Human Spine 253 duties between males and females. changes in physical stress will be due to new repetitive tasks, such as planting and harvesting instead of hunting food, which have' if

general on applied even only intermittently for a short daily time, a higher impact in

this bone mass that just statical fo.... (Lanyon and Rubin, 1984). Biomechanically

labour may be physically more demanding explaining the sometimes-found higher

skeletal robusticity in settled human societies'

Additional environmental factors such as migration patterns as well as changes

the in infectious disease load and its possibly linked nutritional status influence cloud

interpretation of the bony picture as well (Trinkaus, 1981; Jacobs, 1985a; Ruff' 1994;

Holliday, 1996; Holliday, 1997). Furthermore, subclinical microtrauma leading in the

long term to degenerative joint disease will be barely visible initially in the skeletal

records.

To summari ze, the well-known decrease in skeletal robusticity and individual

result of stature in the European Paleolithic-Mesolithic transition period to be rather a

and of the selective forces favouring smaller bodies with reduced metabolic demands

robusticity, \¡/eapon sophistication, no longer favouring taller body stature and bony

postcranial diaphyseal than climate or nutritional stress (Frayer, 1981). The decrease of

to robusticity as seen in early modern Homo as well as in living humans was supposed

1997)' These be due to a decrease of mechanical loading (Ruff et al., 1993; Trinkaus,

parts in findings could be linked to varying susceptibility of the different long bone

(Ruff et aL, 1994)' different periods of ontogeny, such as adolesceîce versus adulthood

are also true for Whether these assumptions on the importance of environmental factors

evaluation' the spinal morphometry would be crucial to know and would need further

255 F. J. Rühti - Osteometric Variation of the Human Spine Alterations of brain and skull morphometry as models for the spinal microevolution

If one discusses changes of spinal morphometry, it is necessary to be aware of

evolutionary trends acting on the other major part of the human nervous system, the

brain, too. At least for the braiñ size such trends have been widely explored.

The evolution of the brain size is supposed to differ from the one of the spinal

cord (Maclarnon, I996a). Relative to body size spinal cord size varies less than brain

size in living species (Maclarnon, 1996b). Since the Late Pleistocene human brain size

seems to have decreased by approximately 10% (Wiercinski, 1979; Henneberg, 1988;

Henneberg and Steyrì, 1993; Ruff et al., 1997). This reduction of absolute brain size

over the past 35,000 years appears to be paralleled by a decrease in average body size

(Ruff et al., 1997). It is assumed that brain size in mammals is a representation of

metabolic rate and not primarily body surface area (Martin, 1981). Brain size may be

related to lean body mass and body height rather than to body weight (Holloway, 1980),

which includes in humans to a highly variable degree the metabolically mostly inert fat

tissue (Henneberg, 1998). How close the relation between metabolic rate and brain size

or neural tissue size in general might be, could be questioned, since its relation seems to

be much more diverse than just a representative of a trade off between gut and brain

(Henneberg, 1998). Nevertheless, the human brain I body size ratio is postulated to be

induced by structural and functional reduction of the gastrointestinal tract (Aiello and

Wheeler, 1995) or as a "structural reduction" of the musculo-skeletal support,

respectively (Henneberg, 1995). A total of approximately 40o/o of the gastrointestinal

and masticatory complex size seem to be lost as a secondary adaptation, which, to

summarize, can be linked to changes in overall body size of about one third (Aiello and

F. J. Rühli - Osteometríc Vartøtion of the Humøn Spine 256 'Wheeler, 1995; Henneberg, 1998). The gut size reduction appears to be related to richer meat-based diets and improved extra-oral food processing, which is supported by

increased mental abilities. Since brain size correlates well with muscle mass (Rogers,

1992), the brain size decrease in the Holocene with its structural body alterations does

not surprise. Brain size and intelligence or mental capacity are weakly or not correlated

at all (WilleÍnan et al., I99l; Henneberg,1992), therefore, the brain size reduction may

be based more on structural reorgarization and increase of neuronal efficiency than just

represent a loss of neuron numbers. The decrease in brain size, with miniaturization of

its neuronal cells, has been explained to be a result of ecosenitive influences in a form of

a decreased meat consumption, not a general decrease in nutritional supply (Wiercinski,

1979). In general, the alterations of brain size in recent human evolution show the

plasticity of the central nervous system in humans and, therefore, raise expectations of

similar trends for the size of the vertebral column.

In general, the size of neural structures might not reflect in a simple evolutionary

way its function and, in particular, the extent of its demand. For example, it is still

debated, if humans, due to the increased demand for motor control and bipedialism,

require greater mass of motor cells and, therefore, show larger neural canal dimensions

than their extant hominoid relatives (Sanders, I99l).In rats, there is apparently a link

between the size of the innervated tissue and the calibre of the cervical nerve roots

(Dunn, l9l2). Furthermore, one has to be aware that the number of somatic afferent and

efferent nerves must not correlate with the body surface area (Fox and Wilczynski,

1986). Differences in various sensory modality systems or density of body surface

innervation, depending on body síze, may account at least partially for such

inconsistencies (Fox and V/ilcz¡mski, 1986). In addition, Agduhr (1917) already found

F. J. Rùhti - Osteometric Varialion of the Human Spine 257 an increase in size of these parts of the spinal cord, which were object of forced training, as shown for growing cats.

The increase of brachycephalisation, another example of microevolution in humans, has been found in Central Europe to be much more coÍrmon nowadays than it was in earliertimes (Henneberg, 1976); however, not all areas intheworld show such an ongoing brachycephalisation trend (Henneberg and Steyn, 1993; Kouchi, 2000). As one possible interpretation of the selective pressure acting on skull form, a differential morbidity of brachycephalic individuals caused by childhood diseases has been mentioned earlier (Henneberg, 1976; Henneberg et al., 1984a). As outlined above, there are some links between spinal morphometry and the occurrence of pathologies, such as lower back pain, but its evolutionary impact appears doubtful. Additionally, climatic influences such as temperature and humidity ecozones have been linked to head form

(Beals, 1972), following the rules of Allen (1877) and Bergmann (1847). In general, this would most likely affect the spinal morphology as well. Finally, nutritional effects

(Lasker, 1946; Wiercinski, 1979; Moishezon-Blank, 1992), migration patterns, parental environmental background or genetic influences, such as exogamy or endogamy, the latter interacting with age and social factors, might reflect on the head shape (Palsson and Schwidetzky, 1973; Billy, l9l5; Kobylinasþ, 1983). Again, it seems reasonable to assume that these factors have at least a partial impact on the evolving spinal morphometry as well.

Microevolution and secular trends of the spine and their possible etiologies

Evolutionary forces can be either of directional or more random-like, non- directional type (Wright, 1968). The first one is usually caused by mutations or natural

F, J, Rühli - Osteometric Variation of the Human Spine 258 selection, whereas the second one generally is influenced by factors such as migration or inbreeding. Both main evolutionary forces may alter the human spinal column. In the present study, the overall changes of life-style and environment seem to suggest a

relaxation of natural selection, a phenomenon already proposed in earlier studies

(Henneberg, 1976; Henneberg et a|.,1978; Stephan and Henneberg, 2001). The second

main evolutionary force could be in the present work migration patterns involving large

parts of central Europe during the covered time span. How far each of the two main

forces contributes to the above-presented alterations of human spinal morphometry is

difficult to assess at this stage. More comparative data would be crucial to improve any

conclusive judgement.

In the present study, various alterations of the spinal morphology with time were

found, which can be classified as microevolutionary or secular trends. A range of

variables changes significantly with historic time period either in linear, cubic,

quadratic, exponential, logarithmic or power function forms. Most of the diameters

show an increase towards more modern times, while some e.g., female femur length,

showed a decrease through time. As shown in Figures 19 and 20, there are in both sexes

consistent alterations of mean values, but there are also changes in standard deviations

of various parts of the spinal column.

The changes in mean, as well as in standard deviations, represent two ways of

relaxation of selective forces. The shift of means in any direction is a representation of a

microevolutionary or secular trend, in the present study presented generally by

increasing values. Therefore, this would be a positive directional selection, similar to the

above-discussed example of trends towards brachycephalisation in Europe. The change

in standard deviations reflects an alteration of the overall variability. Since most of the

F. J. Rühti - Osteometric Variation of the Human Spine 259 changes in standard deViations show an increase as well, this indicates that the diversity of the spinal column increased between the Neolithic I Btotue Age samples and the modern ones in the present study. Again, this part would be a disruptive, non-stabilising relaxation of natural selection. Non-stabilising forces will support the expression of diversity and lead to a higher variability of specific morphological traits'

So far, few reports have addressed secular and microevolutionary trends in the

human spine. Jankauskas (1994) found no significant influence of secular factor on

anatomical spinal landmarks, except for the middle vertebral body breadth of cervical

vertebrae. However, Stefko (1926) described a decrease in spinal height in Russian

,.before the historic samples from 1912" and "from 1923 to 1928-, which might reflect

influence of starvation. Additionally, Tatarek (2001) reported briefly significant

variation of the lumbar neural canal in relation to ancestry of the sample as well as in

relation to geographic origin. Furthermore, Minne et al. (1988) highlighted the impact

of a secular increase in stature in the last century on the spinal morphometry. By

comparing the comparative data of Minne et al. (1988), one sees that there may be a

slight secular trend since the end of the 19* century; see also Figure 27. Nevertheless,

being one has to be aware that the represented samples have different methodical origin,

either cadaveric and osteometric or radiological clinical studies; this may bias the

reported trend.

If one analyses the few historic reports on spinal morphometry available, which

equivocal are comparable in terms of measurements with the present study, one finds

vertebral results. No consistent and clear secular trend is e.g., visible in the L5 ventral

to be an body height in the two sexes; see also Figure 28. While in females there seem

260 F. J. Rühti - Osteometric Variation of the Human Spine

40

o eß g t. o g É t o É al g2 ø ô t + o Eó t o I DÊ o ¡ 2S o G t + a ô of, ó * a to d ¡t I þ ü' t a 21 o9 rt o ð ð * a ô t ! ô 8o * 20 ;g * a ¡ a Ed I à o 13

Aeby Dwight Jacobl and Hurxthal Minne ta"*r, {å76 fSss schmor¡ ls68 I 986 Sludy 1926 2003

Figure 27 Secular trends of male and female spinal dimensions, v/ith vertebrae Th4 - L5 included for earlier studies and Th6 / Th10 I LI I L5 in the present study (Figure modified after Minne et al. (1988), for listed references see there. The data by Hurxthal and Minne are obtained from radiological measurements, with all others resulting from osteometric studies).

F. J. Rühli - Osteometric Variation of lhe Human Spine 262 \ E mm mm :- oa (Ð rù ru N) tu ru cô (¡) o) oJ r\)lutufÞtulÐrÞN)Na r-É oN)Þo)@ ÒN)Þo)@ lñõ+'cno)-.¡æ(oo CD N) I oo o Predmosti 4 Predmosli 3 oG Téviec G ã< Téviec ì CD' il Õ\ ¿ltÄ ì c)<Þõ Þ G CDO (¡ o Ø= Lithuanian 1st 2nd Mill. A.D. Ëi+ Lithuanian 1st / o ,o QH. Fdx 2nd Mill. A.D. o G (DHÊ- q¡- 3.¿ 9¡_ ¿DS o fDg o Early Medieval Polish Þ.ä Early Medieval (D o & E at Polish ct (\ )=' !¡- Èg q¡_ ct 6q) E o -(loã- o Polish, 12th century CL tsC) CL .tl Rural 12th-14th ¿ o- o cent. Polish 9. Rural 12th-l4th century Polish (o (c¡ il o J r_t Urban 14th-18th 9 = ll Polish o (D cent. Polish Urban 14th-18th century o d o (Jl 3 >1 b o (¡) q¡ Þ o x @ 3 o @x + (t, (D!l (D ln ft I Þ) Ð 9 il è (t (¡ Polish o) É Polish ll I l\) Fi t\) o x o @ o) CD x o H è-t \¡ o Or + Recent Europeans E Europeans o o ¿o p io (o@ (o @ -Ø (o x Þ Whites d Italians, American + êl + l\) Ø premenopausal Ì\) @ CD CD co\ Ê- .\¡io American Whites (o Italians, o,NJ o (¡) ) postmenopausal 16.0

! u.o ocft

o 12.0

10000 7500 400 5000 2500 0 FLMl years BP

o

o

35.0

G' IE -o so.o st-

25.0

500 20.0 475 10000 450 7500 5000 425 2500 0 FLMl years BP

(whole sample; p< selected variables in males with significant alterations Figure 2g: with time 0.05; after Bonferroni's correction for multiple comparisons) maximum femur length Ml) and Th 10 sagittal linear regtession Plane.

2& Human Spine F. J. Rühli - Osteometric Variation of the 8.0

7.0 É, I ,.0 o lJ- o 8un oo 4.0

3.0

475 1 0000 450 7500 5000 425 2500 400 0 FLMl years BP

s5.0

o) 50.0 =ro o J ¿s.o a

o 40.0 og

3s.0 475 450 I 0000 7500 425 5000 2500 4oo 0 FLMI years BP

Figure 30: Selected variables in females with significant alterations (whole sample; p< 0.05; after Bonferroni's correction for multiple comparisons) with time before present (years BP) and independent of maximum femur length (FLM1): C3 left cranial intervertebral foramen width (C3IFLCR) and L5 vertebral body transverse diameter (L5M9) shown with linear regression plane.

F. J. Rühli - Osteometric Variation of the Human Spine 265 tries The human body is influenced by a continuously changing environment and

have an impact on to adapt to an energetic optimum. At the same time, these alterations theenvironment.Thisself-amplifl'ingfeedbackbetweenhumansduringevolutionand their living conditions (Bielicki, 1969) will certainly affect the human spinal human biological morphometry. Environment, social otganízation, technology and through human characteristics form a self-amplifuing feedback regulator system

As mentioned above,. evolution as well as in microevolutionary and secular adaptations.

morphological any alterations of natural selection influence the variability of human

act in traits (Henneberg et a1.,1978). Various "cultural" and "non-cultural" mechanisms

such a positive ecological framework (Bielicki, 1969)'

morbidity as In general, natural selection acts through differential mortality and

hard to be replicable in expressed by various levels ofreproductive success, all ofthem

pools are usually slower terms of specific spinal morphometry. Modifications in gene

can be of various forms' as than adaptations to a changing environment. The latter one in Poland and the example of the coincidence of introduction of a feudal social system

(Henneberg 1976)' It seems worth to be the spread of brachycephalisation shows '

further investigated whether similar socio-cultural events would explain spinal

osteometric alterations' be of various Possible etiologies of secular and microevolutionary trends could natural fertility' origin: Decrease of premature mortaiity, birth-planning masking improved medical improved prenatal care, early childhood vaccination programs, from nomadic to settled technology, psychosomatic stresses, physical activity' changes

or the influence of ways of life, dietary changes - such as decreased protein consumption

therefore, higher exchange modern nutrition additives - greater mobility of people and,

266 F. J. Ri)hti - Osteometric Variation of the Human Spine of less related gene pools,'climate, and alterations of growth rate or socio-economic status have been mentioned so far (Lasker, 1946; Beals, 1972; Palsson and

Schwidetzþ,1973; Billy, 1975; Wiercinski, 1979; Bielicki and Welon, 1982; Wurm,

1982; Kobylinasþ, 1983; Jacobs, 1985a; Henneberg, 1992; Moishezon-Blank, 1992;

Ruff, 1994; Henneberg and George, 1995; Henneberg and Steyn, 1995; Henneberg,

1997;Tnnkaus, 1997; Henneberg and Louw, 1998; Hukuda et a1.,2000; Kouchi, 2000).

As one anecdotal example, even the influence of changes in baby sleeping positions as a possible factor of microevolutionary trends of cranial shape has been discussed, but ruled out as etiological factor (Kouchi, 2000). Nevertheless, it would be worth to be further investigated how a change in subadult behaviour actually influences adult spinal morphology.

Furthermore, one spinal variation, the incidence of spina bifrda occulta in various historical and geographical samples (Henneberg and Henneberg, 1999), could be explained by several factors such as the level of fluoride in the drinking water (Gupta et al., 1995), variation between so called civilized versus non-civilized populations

(Post, 1966), better living conditions, improved diet and vitamin 86 and 812 supplementation (Elmazar et al., 1992) or by interbreeding and genetic isolation

(Macchiarelli, 1989). Another anatomical variant of the spine, the occurrence of a foramen transversarium bi-partitum, shows a secular increase mostly between the Late

Roman Period and the Medieval Ages (Susa and Varga, 1981). Furthermore, Porter and

Pavitt (1987) showed a possible influence of juvenile stress on the spinal canal dimension, based on two archaeologic samples. Their study is of high value, because it is a rare attempt to link historic environmental factors, possibly even acting in utero,

F. J. Rühli - Osteometríc Variation of the Human Spine 267 with clinically relevant alterations of the spinal morphology. Some of the factors

influencing the occurrence of spinal variants could be important for the alteration of the

non-pathologic spinal morphology as well.

Hormonal influence.on microevolutionary trends, such as decreased skull size,

has been postulated earlier (Henneberg and Steyn, 1995). A change in a few or even just

in one allele is needed to alter significantly a hormone, its receptors or its physiological

response. Clinical syndromes such as e.g., achondroplasia, which involve the skeletal

morphology, depend on just a single point mutation. Any alteration of hormonal levels

and activities during human evolution seems to be quite likely (Rühli and Henneberg,

2002). Hormones and similar acting substances under genetic or environmental control

have an important influence on growth and functional adaptation of a whole variety of

human tissues. Earlier reports already postulated a hormonal-based microevolution of a

cranial variation as well as possible microevolution of a selected part of the postcranial

skeleton (Rühli and Henneberg,200I; Rühli and Hennebetg,2002; Rühli et a|.,2003),

therefore, this seems quite likely for the vertebral column too.

Nutritional factors have already been related to secular and microevolutionary

trends in humans (Frayer, 1984). A low animal- / high vegetable-protein diet and rice

eating ,ù/as proposed as possible etiology for the altered prevalence of cervical spine

pathologies in historic Japanese samples, rather than genetic or repetitive mechanical

factors in form of the traditional salutative bowing (Hukuda et a1.,2000). The general

intake of proteins, but also alterations in baby feeding practices in form of shortened

breast feeding time and early onset of artificial protein rich diet, were related to changes

in individual stature in various historic time periods in Germany (Wurm, 1982). Milk

F. J. Rühli - Osteometric Variation of the Human Spine 268 protein seems to have the biggest impact on skeletal growth, with animal protein and vegetable protein being of less importance (Wurm, 1982). At last the high stature of the

Medieval samples in the present study could be related to the specific nutritional

conditions in the samples origin.

Various authors addressed the fact that changes in lifestyle such as the transition

from a hunter-gatherer to a more settled agriculturalist way of life cause adaptations in

the postcranial skeleton morphometry (Larsen, 1980; Larsen, 1981; Larsen, 1982;

Bridges, 1989), as already discussed above. Apparently, it is more likely that decreased

mechanical load rather than reduced protein intake has caused the changes in postcranial

skeletal dimensions (Larsen, 1981). Beside socio-cultural and technological ecological

changes, a climatic shift from a Pre-Würm maximum towards the present Inter-Glacial

state has been mentioned too (Jacobs, 1985b). The short-term stature alterations seem to

be rather linked to nutritional influences, whereas long-term effects as for example

changes in body breadth, expressed by bi-iliac breadth, may be genetic adaptations to

influences such as climate (Ruff, 1994). For example, this could explain some of the

differences found in body proportions in Europe between Neandertals and modern

Homo sapiens. Genetic drift and gene flow, due to the lack of genetic group isolation for

the first phenomenon and due to continuous population migration in early European

history, has been ruled out for the found alterations in stature, cranial shape, tooth size

and general robusticity (Frayer, 1984). Furthermore, due to the parallel decrease in tooth

size and tooth variation, "relaxed natural selection" has not been regarded as

accountable for these trends, but rather directional selection has been proposed to be

responsible (Frayer, 1984). However, both, natural selection and the "probable mutation

effect" were suggested to cause the human dental changes (Brace, 1963; Calcagno and

F. J. Rühli - Osteometric Variation of the Human Spine 269 known; this is in Gibson, 1988). The true genetic factors causing such alterations are not particular factual for the spinal morphometry'

various Furthermore, the morphometry of the human spine may be altered by

an unaffected factors such as degenerativè changes, aging or injuries and diseases. In

would not be relevant, vertebral column, as selected in the present study, these factors

above' except for the normal age-related influences, as discussed

was As another etiology, a difference in life style between males and females

changes, as seen in selected suggested to be at least partially responsible for secular

between parts of the postcranium (Rühli et a1.,2003). Such a difference in life style too' To males and females would most likely influence the spinal morphology factors of the altered summarize, as already Frayer (1984) admitted, the underlying

human morphological characteristics are diffrcult to explore.

Additionally, it is difficult to point out how specif,rc factors influence various impact of poor socio- body elements, as can be seen exemplifred by the selective the general living economic conditions in children (Henneberg et al.,1998). Apparently,

the human body, with the environment finds a different response on various parts of

usually to be less trunk length, as a representation of the vertebral column in the living,

parts' such as the long bones dependent on these specific conditions than other body

(Henneberg et al., I 998).

The influence of environmental stress on the spinal morphology has been

acting factors and the highlighted earlier (Porter and Pavitt, 1987). Again, the precise but possibly an early most vulnerable period of the human spine growth are unknown,

in later higher risk involvement of stress factors on the human spine development results

270 F. J. Rühti - Osteometric Variation of the Human Spine of clinical conditions (Porter and Pavitt, 1987). The importance of growth disruption on adult spinal canal dimensions is well known (Clark et a1.,1985). It is notable that most of the canal dimensions of the human spine are acquired intrauterine, which makes them more vulnerable to influencing factors at this early stage of individual development

(Clark et a\.,1985). However, the lumbar spine, for example, shows a greater variability

after birth than the other parts of the human vertebral column (Schultz, 1961).

Therefore, any microevolution of an environmental factor will interfere with the

vertebral column growth to various extents at different times of an individual's life.

The various factors influencing human spinal growth must be taken into account

too (Roaf, 1960), since at least some of them may also be relevant in the present study.

One can differentiate between intrinsic and extrinsic factors, such as infectious or

hormonal influences (Roaf, 1960). For most spinal disorders, it is not well known which

of the altered osseous or soft tissue factors actually is of primary and which is of

secondary nature, and the various elements of the human spine have an independent

growth pattern interacting with each other (Roaf, 1960). One can now assume that

misbalance acting even on just one of these structures could have an influence of the

appearances of all spinal structures. Apparently, every vertebra shows a different growth

property; in general the thoracic and lumbar spine shows an almost exponential growth

during childhood and adolescence (Roaf, 1960). Any growth disturbance of the

vertebral bodies have the highest impact on the surrounding structures of all major

spinal parts (Roaf, 1960). It is not possible to assess the relative impact of intrinsic

embryologic and extrinsic mostly mechanical factors on the growth of the human

lumbar vertebrae (Larsen, 1985). Its own growth pattern and the one of the surrounding

F. J. Rühli - Osteometric Variation of the Human Spine 271 tissues influence the morphologic appearance of the human spine too' For example,

Huizinga et at. (1952) linked the narrowing of the lumbar spinal canal to a possible early growth artest effect. i Other well-known features influencing human skeletal growth are infection and

psychogenic factors. Both are diff,rcult to assess in skeletal remains and their magnitude

depends on the type and timing of onset. Spinal morphometry and general health status'

as expressed by number of specific disease episodes and general practitioner

attendances, are known to partially correlate (Porter et a\.,1987)' People with a narrow

sagittal lumbar spinal canal have e.g., more episodes of childhood infections'

Furthermore, epigenetic and intrauterine environmental influences seem to have a

higher importance than genetic factors in developing the sagittal spinal canal

dimensions (porter et a1.,1987). Enzymatic events, acting between the eight to the 16'b

week in utero, the most size-accelerating period, rather than maternal malnutrition,

(Porter appear to be likely responsible for such spinal morphometric developments e/

at., 1987). However, any spinal growth retardation must not necessarily be linked with

general growth retardation (Porter et a\.,1987). Nevertheless, there seem also to be an

the association between educational performance and spinal morphometry, as shown by

relationship between schoolchildren test scores and lumbar sagittal spinal canal

diameter. Whether this correlation is due to increased sickness-related school absence in

the sample with the naffower canal or whether there is a real link between impaired

(Porter neural canal diameters and early childhood neural development could not be said

et a|.,1987).

272 F. J. Rühti - Osteometric Variation of the Human Spine Based on all these etiological reports on human microevolutionary and secular trends, it is difhcult to end up with a convincing hypothesis to explain the found trends of alterations in spinal morphology in the present study. More research would be necessary to focus specifically on selected factors. It is most likely, however, that spinal dimensions are related to body size while reflecting complex demands of biomechanics

and protection of nerye pathways. While some of these factors seem to be less likely,

such as gene flow, others such as locally different nutrition e.g., in the Medieval Age

samples with apparent tall individual stature, could explain at least some of the

morphologic alterations.

Importance andfunctional implications of osteometric spínal data

The important value of spinal morphometric studies for various research fields

such as anatomy, orthopaedics e.g., for the precise manufacture of surgical implants or

screw insertion depth and direction, biomechanical studies e.g., the use of vertebral

body replica in anthropometric-ergonomic studies or comparison with established

models of animal spines has already been shown (Saillant, 1976; Kikuchi et al., 1977;

Nissan and Gilad, 1984; Nissan and Gilad, 1986; Ziñnck et al., 1987; Ktag et al',

1988; Marchesi e/ al., 1988; Banta et a1.,1989; Misenhimer et a1.,1989; Olsewski e/

al., 1990; Weinstein et al.,1992;Hou et al.,1993; Vaccaro et al.' 1995; Xu et al',1995;

Kothe et al., 1996; Ebrahetm et al., 1997; Karaikovic et al., 1997:' Kandziora et al.,

200I; Mitra et a\.,2002). Macerated spines in particular have an enonnous potential for

the study of their pathologies (Swedborg, 1974) or their normative data and their

variability can be used for assessing developmental pathologies of the spine (Piontek

and Zaborowski, 1973). Surprisingly, there is still an apparent lack of sufficient

F. J. Rühli - Osteometric Variation of the Human Spine 273 clinically relevant osteometric data of the human spine (Krag et al.;1988). Computer- based simulation in biomechanical studies on the human normal and abnormal spine, as done earlier (Schultz et al.,l9J2), would benefit from a databank of normal osteometric reference values too. Additionally, osteometric data are useful since they match well with CT scan data (Be.ry et al., 1987). Furthermore, osteometric reference data of the

spine can be used to detect vertebral crush fractures in individuals who do not show

established patterns of spinal morphometry (Minne et al., 1988). Finally, osteometric

data could also be helpful for studies in forensic anthropology and paleoanthropology

(Jankauskas, 1994). One has to be aware that some osseous dimensions exist, which are

of even higher clinical value e.g., the effective pedicle diameter (Banta et al., 1989),

than the established osteometric measurements. However, this particular measurement

could not be assessed in a non-destructive analysis of historic spines. Nevertheless,

osteometric data gained from historic non-pathologic spines still have their real value

such as e.g., by exploring historic dimensions of spine pathologies.

Spinal morphology has been linked to important clinical pathologies such as

lower back pain, in form of e.g., a link between the circular shape of the vertebral

endplate and the occurrence of a disc herniation (Harrington et a1.,2001), a correlation

between the presence of sacralisation of the most lumbar vertebra and sacral pain

(V/illis, 1924; WllIis, 1929; Philipp, 1932; Gtll and White, 1955) or the size of the

transverse process at L5 and the occurrence of lower lumbar degeneration (MacGibbon

and Farfan, 1979). Genetic or mechanical factors influence the spinal morphology and

may be responsible for the interaction between stature, general muscular and regional fat

build-up and the prevalence of lumbar hemiated discs (Heliövaara, 1987). In skeletal

F. J. Rühli - Osteometric Variation of the Human Spine 274

industrialized modern societies in the present study demonstrate a mild secular alteration of the intervertebral foramen, even without an apparent major shift in culture.

Surprisingly, in the presenl gamples there was also no correlation between osseous intervertebral foramen dimensions and stature or age at death, unlike in previous clinical reports (Humphreys et aL.,1998).

Changes in general bony robusticity, as expressed by femoral robusticity, rather than stature, could at least partially explain any secular alterations of the intervertebral foramen size. This is not the case in the modern samples of the present study, which show an insignificant positive increase in robusticity. A positive increase of robusticity would quite likely oppose a secular enlargement of the mostly bony enclosed foramen space.

The stronger expressed secular trends in intervertebral foramen size in females, in the modern samples of the present study, lack arì evident interpretation and would need further exploration; especially, since in recent samples intervertebral foramen and spinal canal si'ze show mostly no significant sexual dimorphism (Lee et al., 1995;

Ebraheim et a|.,1996).

The results from the present study proclaim a secular na:rowing of the intervertebral foramen diameters, as a possible microevolutionary pre-condition of radiculopatþ or general spinal stenosis, to be unlikely. The mild secular trend of the intervertebral foramen diameters mùy not correlate with alterations in clinical presentation, since earlier studies focusing on possible links between altered spinal neural pathways and clinical symptoms showed inconsistent results (Boden et al.,1990;

Hasegawa et al., 1995; Humphreys et al., 1998), For example, an astonishingly high

F. J. Rühli - Osteometric Variation of the Human Spine 277

Any microevolutionary or secular spinal trend may represent so called "relaxed natural selection", which is particularly visible in developed countries and may decrease the ability of humans to su1ive and reproduce without medico-technological help

(Stephan and Henneberg, 2001). The selection forces acting particularly on the human

spinal column are still mostly unknown. For example, humans, despite having a much

larger relative brain weight, do not have a bigger spinal cord weight in comparison with

other primate and mammal species (Maclarnon, 1996b). Therefore, one can assume that

the selective powers influencing the spinal morphology must be different from the ones

interfering with the other central nerve system part, the human brain. Possible etiologies

of the findings in the present study may be, as pointed out in earlier microevolutionary

studies (Wiercinski, 1979; Wurm, 1982; Henneberg and George, 1995; Rothschild and

Rothschild, 1996; Henneberg and Hennebetg, 1999; Hukuda et al., 2000), based on

genetic e.g., changing allele frequencies, or environmental influences e.g', nutrition. In

general, the microevolutionary and secular trends in the present sample show that there

is ongoing influence, mostly balanced between environmental and genetic factors,

which acts on the human spinal column.

The challenging results, as presented above, will hopefully stimulate the debate,

which assesses spinal morphology changes by using a historic perspective (Clark et al.,

1985; Porter and Pavitt, 1987; Jankauskas, 1992: 1994; Henneberg and Henneberg,

1999; Hukuda et a1.,2000; Boszczyk et a1.,2001; Tatarek, 2001; Rühli et al',2002)'It

builds a bridge between anthropological approaches and clinical research, and it should

help to improve our still limited knowledge on the ongoing evolution and the

osteometric variation of the human vertebral column'

279 F. J. Rühli - Osteometric Variation of the Human Spine

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F. J. Rühlí - Osleotnetric Variation of the Human Spine 333 Appendix

.. 335 1. List of abbreviations for measurements used 337 2. Complete original data

Complete statistics of variables of majortime groups ""' """349

samples (means of modern 4. sexual dimorphism of all variables in modern

samples, p ercentage-difference, t-values)

(means, t-values) 375 5. Side differences of variabies in modem samples "" 376 6. Correlations of variables with individual age in modern samples ' " "' samples 380 7. correlations of variables with major age groups in non-modern " "' major time g. Correlations of variables with major age groups separately for ..384 groups 396 9 Inter-correlations of variables ..420 10 Signifrcant microevolutionary regressions ' ' "

groups 422 11. ANOVA of variables with major time

group I versus 3 (F- t2 Alterations of standard deviation of variables with time values) " 430

with time before present and selected 13. partiai correlation coefficients of variables

with the selected long bone measurements (variables with signihcant correlation

434 long bone measurements only; whole sample) ""

14. PrinciPal comPonent analYsis 439 15 Abstract on PhD data (Rühli et al',2002)

334 F. J. Rühti - Osteometric Variation of the Human Spine 1. List of abbreviations for measurements used Abbreviation Variable

BP year of birth before 2000 A.D. Agegroup Adult (Agegroupl); Matur (2), Senil (3) c3M2 C3-dorsal vertebral body height C3M1 C3 ventral vertebral body height c3M6 C3 sagittal diameter vertebral body c3M9 C3 transverse diameter vertebral body C3PHI C3 left pedicle height C3PHr C3 right pedicle height C3M1O C3 sagittal diameter spinal canal C3M11 C3 transverse diameter spinal canal CSSPL C3 spinous process length C3TPW C3 transverse process width C3lFlcr C3 left cranial intervertebral foramen width C3lFlca C3 left caudal intervertebral foramen width C3lFrcr C3 right cranial intervertebral foramen width C3lFrca C3 right caudal intervertebral foramen width c7lul2 C7 dorsal vertebral body height c7M1 C7 ventral vertebral body height c7M6 C7 sagittal diameter vertebral body c7M9 C7 transverse diameter vertebral body CTPHI C7 left pedicle height CTPHr C7 right pedicle height C7M1O C7 sagittal diameter spinal canal C7M11 C7 transverse diameter spinal canal CTSPL C7 spinous process length CTTPW C7 transverse process width CTlFlcr C7 left cranial intervertebral foramen width CTlFlca C7 left caudal intervertebral foramen width CTlFrcr C7 right cranial intervertebral foramen width CTlFrca C7 right caudal intervertebral foramen width T1M2 Th1 dorsal vertebral body height T1M1 Th1 ventralvertebral body height T1 M6 Th1 sagittal diameter vertebral body T1 M9 Th1 transverse diameter vertebral body T1 PHI Th1 left pedicle height TITPHr Th1 righ pedicle height T1M1O Th1 sagittal diameter spinal canal T1M11 Th1 transverse diameter spinal canal TlSPL Th1 spinous process length TlTPW Th1 transverse process width Tl lFlcr Th1 left cranial intervertebral foramen width Tl lFlca Th1 left caudal intervertebral foramen width Tl lFrcr Th1 right cranial intervertebral foramen width Tl lFrca Th1 right caudal intervertebral foramen width T6M2 Th6 dorsal vertebral body height T6M1 Th6 ventral vertebral body height T6M6 Th6 sagittal diameter vertebral body T6M9 ThG transverse diameter vertebral body T6PHI Th6 left pedicle height T6PHr Th6 right pedicle height T6M1O Th6 sagittal diameter spinal canal T6M11 Th6 transverse diameter spinal canal

F. J. Rühli - Osteometríc Variation of the Human Spine 335 T65PL Th6 spinous Process length T6TPW Th6 transverse Process width width T6lFlca Th6 n en width T6lFrca Th6 T1OM2 Th1 body height T1OM1 Th10 ventral vertebral vertebral body T1OM6 Th10 sagittal diameter vertebral body T1OMg Th10 transverse diameter TlOPHI Th10 left Pedicle height TlOPHT ThlO right Pedicle height spinal canal T10M10 Th10 sagittal diameter spinal canal T10M11 Th10 transverse diameter length TlOSPL Th10 sPinous Process width TlOTPW Th10 transverse Process foramen width Tl0lFlca Th10 left caudal intervertebral foramen width Tl0lFrca Th10 right caudal intervertebral bodY height L1M2 L1 dorsal vertebral bodY height L1M1 L1 ventralvertebral vertebral body L1M6 L1 sagittal diameter vertebral body L1M9 L1 transverse diameter LlPHI L1 left Pedicle height height Ll PHT L1 right Pedicle sPinal canal L1M1O L1 sagittal diameter spinal canal L1M11 L1 transverse diameter LlSPL L1 sPinous Process length wdth LlTPW L1 transverse Process foramen width Ll lFlcr L1 left cranial intervertebral foramen width Ll lFlca L1 left caudal intervertebral foramen width Ll lFrcr L1 right cranial intervertebral foramen width Ll lFrca It rilnt caudal intervertebral height L5M2 L5 dorsal vertebral bodY bodY height L5M1 L5 ventral vertebral vertebral body L5M6 L5 sagittal diameter vertebral body L5M9 L5 transverse diameter L5PHI L5 left Pedicle height L5PHr L5 right Pedicle height sPinal canal L5M1O L5 sagittal diameter spinal canal L5M11 L5 transverse diameter length L5SPL L5 sPinous Processus width L5TPW L5 transverse Process foramen width L5lFlcr L5 left cranial intervertebral foramen width L5lFlca L5 left caudal intervertebral foramen width L5lFrcr L5 right cranial intervertebral width L5lFrca L5 riint caudal intervertebralforamen magnum FMM16 sagittal diameter foramen magnum FMMT transverse diameter foramen HLMl maximum humerus length erence HCMT humerus m inimal circumf FHM18 femoral head width FLMl maximum femur length FCMS mid{em ur circumf erence BIWM2 bi-iliac width 336 of the Human Spine F. J. Rähti - Osteometric Variation 2. Complete original data (*: not specihed for reasons of anonymity) : ..-.""..Ë

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.o L1M11 LlSPL LlIFLCR L1 IFLCA LlIFRCR L1 ¡,¡19 L1 PHL L1 PHR L1M1O 19 I 2A N Val¡d 29 29 29 70 50 776 12 17 8.25 38 73 '15 53 16 1s 17 16 22.8O 29 60 12 20 800 '15 50 16 50 17 30 22 70 29 90 70 10 770 : M€d¡an 39.20 11 904 800 5 goa 1650ã 17 50 22 70 29 90 62 404 770 Mode 35 f6 80 U 108 139 159 283 379 1 19 136 \- Std O€v¡ation 2to 18 80 25.00 62 40 ,l 80 910 690 M¡n¡lìm 34 30 13 00 13 80 14 10

L5PHL L5PHR L5M1O LsMl t L5SPL L1 L5M2 L5M1 L5M6 L5M9 31 30 27 27 14 Valid 26 31 '16.26 40 26 72 28 24 32 57 ¡704 14 17 l4 69 25 Mean 12 08 24 27 26 15 90 32 10 47 65 14 10 14 80 15 80 25 65 Median 12 15 24 30 27 'Í 80 12 6oa '15 00 24 ßè 22 502 13 20 24 30 27 50 32.20 48.80 215 244 341 177 194 1-81 204 3 t9 Std Deviat¡on 129 204 22 50 AO 39 30 10 80 11 40 13 00 21 50 Min¡mm 930 '19 20 2s 80 24

OJ(Jr HLMl L5IFRCR LsIFRCA FMM16 FMMT LsTPW L5 FLCB LSIFLCA 24 32 26 10 I 30 2A à Valid 12 N 313 13 62 81 46 04 59e 961 36 15 29 94 83 31 5.84 963 Mebn 3lo oo 62 50 46 70 605 980 36 30 29 80 I 86 55 5.75 too0 Med¡an 305 ooa 62 ooa 16 70 530 980 37 00 29 80 o 45 80ã 4.80å 10 20 Mod€ 181 '14-08 413 89 173 257 s 17 15 .99 1.54 Sld- Dev¡ation 53m 38 80 580 32 80 27 00 291 00 4.30 640 1.20 Minimm 45 80 ô ì o q FLMl FCMS 34 12 N Val¡d 437 85 86 21 Mêan 50 435 00 87 00 273 Med¡an 421 00à 89 00 259 ooa Mode 20 93 581 10.77 tr Std DÉv¡at¡on 76 00 259 00 Minirum 391 00

a. Mult¡ple @des exist. The smllesl valæ is shown

(J) (Jl l-) S Females - Neolithic / Bronze Age à

AGEGROUP caN42 c3M1 c3f,46 c3M9 C3PHL C3PHR c3t\410 c3M1 1 C3SPL I o N Val¡d 54 39 37 34 35 27 14 'I 39 1217 12 1s 14 97 't9 38 621 609 14 65 222e 13 00 .|00 o Med¡an 12 20 12 40 14 80 19 30 610 590 '14 70 22 45 12.S0

Mode 100 12 20 12 60 ttzou 20 80 5 50' 580 is 20' z1 oo" 11 20

Cl Std Dev¡at¡on 63 91 113 234 94 7A 103 1 18 237 ^\\ Min¡dm 100 10.20 10 00 12 50 14 90 420 480 12 10 19 20 940 Þ o

C3TPW C3I FLCR C3IFLCA C3IFRCR C3IFRCA c7M2 c7¡,t1 c7M6 c7M9 CTPHL N Val¡d 17 g4 36 36 34 35 33 ò 48 68 573 702 604 7 -34 13 32 12 95 15,O9 25 06 654 b \- M€dian 48 80 570 730 820 710 13-30 13 00 f5 05 25 20 ô50 Mode ¿e ao" s zo" 7.gou 3 Boa s40 13 30 ts zo" te so' 680

Std Devial¡on 244 131 161 'I 51 171 97 102 1 t9 212 88

M¡n¡mm u2c 360 380 330 390 1 l.s0 10 60 12 60 t9 60 480

CTPHR c7M10 c7M1 1 CTSPL CTTPW CTIFLCR CTIFLCA CTIFBCR CTIFBCA T,1M2

N Val¡d 20 10 34 31 29

Mean 636 1,1 0'l 23 10 27 32 4E 24 620 914 422 838 15 s6

Mediah 6_50 14 30 28 10 46 15 620 910 615 920 15 30

Mode s go" ts go" zz ao' 27 ao" g4 so' 640 900 580 10 00 '15 10

Sld D€vialioh 92 100 149 4-14 11 64 118 111 1.21

Minirum 490 11 40 18 30 15 90 34,30 440 580 450 650 13 30

g) (Jr (]) \r

S TIMI T1M5 T1M9 T1PHL T1PHR T'IM'10 TlM1l TISPL ÍITPW T1IFLCR à 17 27 N Val¡d 2A 2A 29 30 27 15

Mêan 14 68 '!554 26 37 830 836 14.5 1 19 91 27 52 68 28 625 I 69 50 620 o Med¡an 14 50 15 65 26 40 830 840 14 10 19.80 27 70 13 904 l5 80 26 20à 7 60â I ooa 1120 19 50 22 002 68 80 580 ñ Modê 72 o Std- Oeviat¡on 119 132 200 107 I 16 92 I 3,1 308 524 60 22 00 57æ 480 : M¡nirum 12 40 12 80 23 20 550 580 12 10 17 Õ ì Þ o

.o T6M1O T1 IFLCA T1 IFRCB T1 IFBCA T6M2 T6M1 T6M6 T6M9 T6PHL T6PHB 2A 29 29 28 N 27 25

21 6'1 24 55 10 48 1032 15 62 Mean 996 628 985 18.81 17 7l '17 60 24.50 10 50 10 40 15 60 Med¡an 980 610 980 18 60 80 21 20 1oa 23 20à 11 20 10 soa f4 6oa Mode 980 610 10 60 fl.70ã 16oOã b 't.33 168 96 .88 96 \- Std Dêv¡alion 122 98 134 123 188 M¡nillla 690 4,30 640 15 90 15 70 18 00 22 10 890 840 14 30

38 36 N Val¡d 29 14 24 24 25 't6 'It 25 34 29 92 13 A2 Môan 16 36 76 5A77 11 65 03 21 68 20.71 20 60 25 50 29 70 f3 95 Med¡an 16 00 15 90 58 80 1t 45 11 20 21 60 '15 goa 50 20 204 24 604 30 20 11 gOã Mode 80 I 004 47 60ã 1320 9 21 154 Í-93 254 1.3€ Sld Dêv¡ation 1.68 5,29 499 184 180 175 24 AO 10 80 Minimm t3 30 900 47.60 670 8.20 18 60 17 80 20 90 57 00

(}) (Jt 'Þ 5 S T1 OPHR T10M10 T1 0M1 1 Tl OTPW 11 0tFLoA TlOIFRCA Lt[¡2 L'IMl LlM6

N Valid 38 37 s6 26 37 41 39

Mean 1404 1528 1679 23 30 s2 57 11 38 11 15 26 68 24 99 27 45 I Mêd¡an 14 30 15 40 16 70 23 20 o 52 70 11 20 11 40 26 60 21 AO 21 50 Mode 14 80 14.604 1 6 20 22 604 47 60ã 12 00 '!2 60 24 40 27 30_ o Sld Dev¡ation 1 43 1.39 1 54 430 501 167 138 204 188 207

Min¡Ém 10 90 10 80 13 50 18 20 44 40 790 880 23 40 21 30 22 90 37 70 ìÞ- Þ a -ô LlMIO 11Ml1 L1 SPL L.l IFLCR L1 IFLCA L1 IFRCR

41 39 41 21 19 34 39 30

Mean 35 08 14 40 14 70 21 99 25 93 62 06 E.59 12 A3 I10

Median 35 10 14 40 15 00 17 20 22 10 2570 64 00 860 12 90 s05 '16,80 Mode g2.aoa 14 10 14 1oa 20 goa 27 20 59 40â 850 1370 I 3oa .\]-b Sld Devialion 292 1 23 138 'I 22 155 330 7_01 1.15 155 112

Minimm 2920 1 t 10 11 80 '15 10 18 20 18 80 44 20 600 910 6_80 43 10 17 20 18 00 19 80

L1 IFBCA L5M2 L5M1 L5M6 L5M9 LsPHL L5PHB L5M1O LsMl l LsSPL N Valid 38 42 44 44 46 41 12 35 40

Mean '12 61 23 4A 26 67 31 29 14 77 '!3 07 13 62 't6 40 25 38

Med¡eh t2 60 23 40 26 45 31 50 44 60 13 00 13 45 16 30 25 35 22 00

Mode íi 8oa 2s7oã 2440¿ 31 0oâ 4570â f 204 12 40 17 20 26 60 20 704 Std Dev¡ation 1 sl 224 254 2ô6 311 147 177 238 275 32l

Min¡rum 10 10 17 70 21 70 23 10 37 30 9-50 10 30 11 90 19 80 17 90 1560 323î 35 90

O) Cn Ctr I LsTPW LsIFLCR LsIFLCA LsIFRCR LsIFRCA FMI\416 FMMT HLM1 I-ICM7 FHBM18

N VaId 39 39 40 12 13 38 13 52

Mean 75 13 6.16 974 625 10 08 35-55 29 36 291 45 57 47 41 30

I o Median a1 25 610 9.80 610 1025 34 95 29 20 288 00 57 00 41 10 Modê 81 20å 610 980 6 1oa 7ao 38 40 25 004 283 004 55 004 40 20 ß o Sld D€viation 1672 84 142 ?1 188 274 300 17.29 414 234 Minìm 40 00 450 5 t0 4-80 830 3f so 25 00 260 00 48 00 36 s0 I o

a

FLMl FCMs 8twM2

N Val¡d 40 45 l9

M€an 409 05 76.16 259 95

Med¡an 402 00 76 00 260 00

Mode 397 00 75 00 267 00 b \- Std Oev¡al¡on 23 37 546 17 17 M¡n¡ruh 381 00 65_00 234 00

a Mult¡pl€ mdês ex¡st. Thê Eßllest valuê ls shown

(JJ (JI o\

a S Males - Medieval Ages >U

CASPL c3M9 C3PHL C3PHR c3M10 cxlMl t AGEGROUP c3M2 c3M1 c3M6 oI o ì Cl È-ì Þ a

ß CTPHL C3IFBCA c7lu6 CsTPW CAIFLCB

Valid ò 775 194 t4 08 17. 26 38 Mean 54 93 30 26 45 05 U 640 620 7 80 15 00 14 20 17 \- M€dian 54 10 14 ao" 1s ao" a0 25 aô 690 57 20 680 .80 5 30' 780 s5 99 35 141 'I 01 125 93 2A Sld Dev¡al¡on 407 21 20 530 520 110 420 12 60 10 20 13 90 Mihilllfr 47.30 110

CTIFHCB CTIFRCA c7Mt0 c7M1 I CTSPL CTIFLCR 64 61 65 64 61 68 N 63 616 988 628 s85 17.25 14 94 25 11 29 04 68 37 Mean 708 620 980 620 985 17 30 14 80 25 25 29 20 72 AO Mediãn 700 6.50 g oo' 6-20 10 40 16 80 15 10 2g 20' ze 5o" og ro" Mod€ 650 76 121 130 221 410 12 93 90 123 Sld Dev¡alion 79 130 480 720 't3 60 17 10 '!7 80 39 90 430 770 Min¡dm 520 12 30

(Jro) N ! ã T1 PHL T1 [¡1 0 T1M11 TITPW N Valid 70 68 70 66 67 61 62 38 52 65 16 11 17 33 28 33 I3t 935 15 44 22 29 31 25 I 7A 40 642 Med¡an 16 00 17 40 28 35 930 920 o l5 40 22.OO 32 00 77_40 620 Mode 15 8oa 16 50 27 loa 930 20â I t5 80 21 80 32 oOã 79 204 6 104 Sld. Dev¡al¡on 146 139 293 1 18 117 102 192 430 6-65 ß 96 Min¡lìM 12 50 12 90 22 A0 680 700 \ 13 60 f7 60 20 70 50.20 460 ì

o

11 IFLCA TlIFRCA T6t!12 T6t\41 T6M9 T6PHR Val¡d 63 61 64 63 60 62 59 6t 51 Mean 10 0s 623 10 13 21 19 19 03 28 12 12 02 12 22 16 12 Median '10 10 00 630 15 2't 40 18 90 28 00 ò 2A 20 12 00 1220 16 10 Mode tl50 6 5oa 6 80å 19004 1980 b 26 80 26 20 11 80 '12 80 17 60 Sld Dev¡al¡on 141 86 12l '1 \- 58 137 215 208 117 1.24 '1.08 Minirum 680 4s0 750 17 30 15 tO 21 60 22 00 880 930 l3 80 31 0n

T6SPL T6IFLCA T10M2 T10¡.,t6 TlOPHL N Val¡d 60 14 30 4ô 46 71 70 69 70 67 Mean 17 19 19.39 1 1.66 '11 36 24 tO 22 45 30 29 34 39 15 55 Med¡an 17 25 19.00 65 05 '1t 50 50 lt 24 20 22 25 30 60 34 80 15 30 Mode 16 10 io 30â 61 8oa 11 40 11 S0 22 50 22 00 28 8oa 34 80 14 60 Std Devial¡on 145 621 604 2.O4 169 159 158 271 3-f6 130 Min¡dm 14 20 10 30 48 00 750 790 19 40 18 30 25 10 21 20 'l.2 a7

(JrO) co

a T1 OIFLCA T1 OIFBCA Ll M2 L1 M1 L1 M6 : TIOPHB T10[.,110 T10M1 l TlOSPL TlOTPW 77 57 24 40 60 a2 N Valid 67 64 25 94 3t 66 18 54 26 59 60 56 f2 31 11 86 28 06 Mêan 1537 16 23 26 20 31 80 I 18 30 60 20 12 55 12 00 27 95 Medìan 16 00 o 00 27 204 27 10 30 10- S0 18 20 24 20 5i 4oa 11 204 12 ¡,tode 14 8oà 15 '1 276 173 493 588 f75 167 69 204 o Std- Deviat¡on 130 1.32 25 20 '15 48 00 870 830 24 20 21-90 ì Minidm 12 60 13 70 20 1270 ì o

-o LITPW LIIFLCR L1IFLCA LlIFRCR M9 Ll PHL PHR L1M1O 11M11 LISPL 71 66 68 7A 69 74 27 N 81 77 29 90 72 67 823 1270 830 40 49 15 44 15 76 17 91 23 90 830 12 80 840 't5 15 70 '17 80 23 85 29 40 72 00 Median 41 10 30 72 00 830 11 80- 840 ooa 14 60â 17 804 23 10 28 10 Mode 39 80 14 149 tol aâ 131 138 450 11 01 105 Std Deviatìon 321 130 \ 980 610 'i3 20 14 60 20 óo 18 50 37 80 620 M¡n¡mm 30 20 12.10

Ls[¡1 1 LSSPL L5M1 L5M6 L5M9 L5PHL LsPHR L5M1O 30 69 71 59 64 N val¡d 66 73 71 26 52 2174 28 66 33 A2 48 05 14 1? 14 70 t6 78 Mean 12 64 2474 60 16 50 26 80 25.00 90 2A 70 33 70 48 60 14 00 14 M€d¡an 24 p 14 80 15 50 23 80¿ 23 80_ 8oa 29 20 32.10 48 8oa 20à Mode 1t 80 23 394 263 324 449 186 205 212 Std Dev¡al¡on 157 204 '17 60 25 60 36 70 11 10 10 90 12 40 21 10 Minidm 960 21 20 21 90

OJ \oCn LsTPW LSIFLCR ! L5IFLCA LsIFBCR L5IFRCA FMMl6 FMMT HLMl l-tcM7 FH8M18 N Val¡d 2ø 67 59 67 58 18 1? 66 85 80 83 75 5-88 998 605 990 37 23 32 19 934 26 65.93 48 99 Median 89 90 5.80 I 980 620 985 37.00 31 50 331 00 65 00 48 80

o Mode 96 00 500 7 ooa 620 980 35.704 30 90 328 00 65 00 48 00 Std Dev¡al¡on o 18 79 87 185 168 216 255 '17 93 4_1a 303 14.80 M¡n¡dm 4.00 640 400 640 32 20 27 90 294 00 57 00 11.20 ì 1ôr 1ã ñ ìÀ\

FLMl FCM8 BIWM2 N Val¡d 68 83 27

Mean 464 40 90-90 2a2 15 Median 464 00 91.00 283.00

ò i,lode ,155 oo aS ooå 280 OOâ \-u Std. Dev¡at¡on 27 66 6 43 17,49 Mìnìmm 376 00 75 00 230 00

a. Multiple mdes sx¡sl- Th€ smllest vâlue is shoM

(, o\ S Females - Medieval Ages

AGEGROUP c3M2 c3M1 csM6 c3M9 C3PHL C3PHR c3Mr0 c3M1 1 C3SPL I o Val¡d 81 63 64 61 61 63 63 50 6.1 22 Mêan 164 12 43 12 27 14 67 18 13 s92 602 14 A1 23 33 1A 12

o Med¡an '| o0 12 50 12 20 14 60 17 90 580 600 15 05 23 20 13 15

80 12 80 zo" '18 00 520 630 't5 60 zg so" t so" ì Mod€ 100 12 l¿ C) Sld Oev¡ation 110 120 184 a7 124 1-33 171 ì Miniflm 100 950 910 12 10 14 60 130 ,l 80 11 70 20 t0 10 10

a .Ò

C3TPW C3IFLCR C3I FLCA C3IFRCA c7M2 c7M1 c7M6 c7t\¡s CTPHL

N 36 61 61 6l 65 66 65 64 64

Mean 49 70 664 a2l 649 8ls t3 75 12 87 15 67 24 A5 674 tr Med¡an 49.35 650 850 670 830 13 80 12 90 15 70 24 SO 675 Mode 42 so" s ¿0" z to^ 680 z oou tz oo' t 1.a0" 15 80 z¿ zo' o zo"

Std Dev¡at¡on 384 1.05 160 108 1.44 108 112 140 95 Minimm 43 20 380 480 420 470 10 80 10 50 12 80 20 80 420

cTpHR c7M1o C7¡¡11 CTSPL CTTPW CTIFLCR CTIFLCA CTIFRCR CTIFRCA T1M2

N Valid 68 60 63 35 17 58 57 59 69 666 14 45 24 57 25 59 58 86 632 971 639 957 15 71

Med¡an 660 14 50 21 60 26 20 65 60 630 980 6.30 970 15 60

Mode s eo' 14 60 25 80 z¿ oo' zl oou 680 980 610 980 1540" 'I Std Dev¡al¡on 79 ,{1 170 291 17 35 95 128 85 127 122

M¡nimm 490 10 60 20 60 19 60 2'f .30 120 650 420 630 13 20

(¡) o\ }1 S T1 M1 T1M6 T1M9 T'I PHL 11 PHR T1MlO T1M1I TlSPL TlIPW T1 IFLCR N Valid 68 64 66 63 69 56 65 25 61 Mean 't4 56 15 83 a47 839 14 90 21 52 26 69 71 08 e34 I Med¡an 14 60 15 85 26 35 860 o 830 11 75 21 20 26 90 71 40 630 Modê 14.OOa 15 3oa 26 00 860 780 t4 50 20 50 22 604 68 soa 620 o Sld Dev¡al¡on 1.09 137 261 1 10 101 132 172 364 514 80 ß M¡nirum 1210 12 80 21 40 5S0 'It ì 580 80 18 40 '19 40 58 20 450 o \ È o q T1 IFLCA TlIFBCR TlIFRCA T6M2 16M1 T6M6 T6N,,IS T6PHL T6PHR T6t\4r0 N 59 65 60 o/ õb 65 67 62

Mean 10.27 630 10 09 'I 't7.39 S 05 23.17 24 95 10 27 10 49 t5 86 Med¡an 't0 s0 640 35 t0 19 00 17.35 23 60 24 90 10 15 10 50 15 85 Mode I 504 6 50À 11 30 18 soa i6 90ã 24 00 22 AOa 980 10 80 16 40 b ,Ù- Std Dev¡al¡on 159 84 146 125 124 2.08 80 122 Minidm 650 440 640 15 't810 G 70 14 s0 21.80 810 8AO 13 30 13 50 o

T6M1 1 T6SPL T6TPW T6IFRCA T10[.,t2 Tt0Ml T1OM6 TlOMg T1 OPHL N Val¡d 62 18 41 50 47 67 66 67 68 65 Mean 16 49 14 81 59 67 1217 12 06 21 67 20 a2 26.39 30 55 13 85 Med¡an '16 40 14 95 58.80 1225 12 00 21 AO 2l 05 26 30 30 75 13 90 Mode 15 20ð 6 8oa 57 20 11 00- l3 00 21 90 21 20 27 50 28 80 14 00 Sld Dev¡alion 144 453 596 158 1.45 157 167 219 122 Minimm f3 00 680 45 50 850 900 16 40 16 00 21 30 25 20 '10 50 20 Áo 15 20

o\OJ l.J þ S TÍOPHB T1OM1O T1OM1l T1OSPL T1OIPW T1OIFLCA T1OIFRCA L'IM2 L1[41 L1N¡6

N Valid 65 63 41 55 71 70 66

Meân 13 8,1 '15.61 17& 23 65 51 A7 12 59 12 09 26 06 24 59 27 53 oI Median 13 80 15 65 17 40 24 45 55 00 12 60 12 00 26 20 24 50 27 40 Mode 13 80 '15 90 17 30ã 24 AO' 53 6oa 13 20 11 304 26 50 24 504 26 20

o Sld. Devial¡on 109 141 132 2.88 525 140 161 1.67 181 234

M¡n¡dm 't0 20 1't 90 1.1 50 15 10 42 10 870 7ao 22 00 19 20 21 70

Ò s

a .a L1M9 L1PHL L1PHB LIM1O 11M11 L1SPL L1TPW L1IFLCR L1IFLCA L1IFRCR

N Valid 6S 66 71 58 69 58 53 57

35 76 14 25 '14 39 't7 73 22 62 25 33 64 09 471 1298 8s4

Med¡an 36 00 14 10 14 50 17,70 22 50 25 25 65 60 850 12 80 860 =s Mode 38 40 15 10 1800å 21 10 14 8oa 36 2oa 850 1220 I 30â lâ \ Sld D€vialion 299 123 140 147 175 386 845 117 136 114 Min¡dm 26 60 1140 to 70 15 00 18 00 14 80 36.20 620 10 00 630

L1 IFBCA L5M2 LsN.,t1 L5t\,ll6 L5t\.19 L5PHL L5PHR LsM1O L5M1 1 L5SPL

N Valid 56 68 62 62 67 63 6,1 57 61 24

Mean 12 89 23 42 26 85 3t 21 44 23 1277 13-62 16 94 26 16 24 20

Median 13 00 23 85 2670 31 30 43 20 12 80 13 25 l6_70 25 90 24 00

Mode 12 10 25 30 25 50ã 32 2oa 41 8oâ 12 204 12 AO 15 204 25 30â 25 ooa

Sld Deviar¡on 136 1,99 265 273 1A7 167 214 253 372 Minimm 10 00 19.00 21 50 24 10 35 00 860 950 12 40 21 20 17 70

(¡) o\ o) -{t

S LsTPW L5IFLCR L5IFLCA LsIFRCR L5IFRCA FMM,I6 FMMT HLM1 FICMT FHBM18 >ù N Valid 31 64 59 6.1 17 17 54 74

Mean 76.99 627 10-60 646 10.4,1 35 90 29 f8 303 89 56 57 43 26 I Median 81 10 630 10 50 630 10 60 35 80 28 80 305 00 56 00 43 20 O Mod€ 87 60 6.80 10 40 6 2oa 11 80 36 00 28 20 292 00 55 00 13 20 a Sld Oêv¡at¡on 15 01 100 1ô1 1 '10 171 228 186 19 11 356 ì M¡n¡llffi 11 10 420 720 4.30 580 33 ÍO 26 20 261 00 49 00 38 00 o ì Þ a

FLM1 FCM8 BIWM2

N Val¡d 64 33

Mean 429.63 80-60 271 79 I Med¡an 435 50 80 00 270 00 h i,lod€ 112 00 79 00 258 ooa b \- Std Oev¡at¡on 24 AA 509 13 44 ô M¡n¡dm 379 00 70 00 24s 00

a Muh¡p¡e mdes e{sl Thê 6Ellesl valuê ¡s shown

(¡) o\È i Males - Modern

AGEGROUP C3[I2 caMl c3M6 csMs C3PHL CSPHR Csl\.,110 c3¡¡11 C3SPL

I 11 N Val¡d 41 38 36 37 34 o 16 95 Mean 202 14 57 14 15 16 20 19 27 726 15 93 24 54 't610 60 16 80 Mêd¡an 200 14 65 14 00 19 40 700 710 15 90 21 '19 go' 24 60 G Mode z oou tg 20" i4 oo" 14 oo" 80 690 700 ts tzzo^ 'I '! 169 296 ì Std Dev¡al¡on a2 136 01 1,49 18'1 1.11 103 45 13 40 '16 70 550 540 13 10 19 90 't2 70 s\ Minirum 1.O0 1 0.50 11 E0

-o

c7M9 CTPHL G3TPW CsIFLCR C3IFLCA C3IFRCA c7M2 C7M1 c7M6 35 38 36 36 N 37 34

56 10 682 ô49 420 15 15 1372 17 65 26 55 754 a- 650 830 15 20 13 70 26 50 \ M€dian 55 70 7.00 820 20 16 50_ z4 so' 640 Mode sg oo' 7 oo' 680 580 830 rs ¿0" 13 94 Sld Devialioñ 376 86 1.33 94 130 124 145 173 620 M¡n¡ruh 47 aO ¡t 80 530 420 4-50 12 20 10 10 l4 90 2220

CTPHÊ C7M1O c7¡,,t11 CTSPL CTTPW CTIFLCB CTIFLCA CTIFRCR CTIFRCA T1M2 35 40 N Valid 38 38 38 7 35 10 05 655 10 10 17 29 Mean 15.t0 26 14 31 53 66 21 630 '10 00 660 10 t0 17 05 Mêdian 750 15.30 26 30 32 20 70 30 650 00 680 g oo" 16 80 Mode 880 f5 50 26 40 zg to" as aou 700 11 73 1 18 141 Sld Dev¡al¡on 97 153 r73 430 18 40 93 149 510 750 14 00 Min¡dm 590 10 60 2270 23.1 0 25 80 480 650

(¡) o\ (Jr S TlMIO T1M11 TlSPL 11 I FLCR T1M1 TI M9 T1 PHL 20 35 40 39 38 ¿0 N Val¡d 38 't5 79 07 656 931 912 78 23.26 33 24 16 03 17 76 28.93 M€an 665 1ô 10 23 20 33 60 78 30 I 17 40 2A 70 955 920 Med¡an 15 85 16 10 21 soa 30 1oa 82 90 7so 17 10 25 60â 780 980 Mode 16 30 ß 121 178 357 509 101 197 27A 126 124 o Std D€v¡at¡on 129 19 50 23-90 69 80 4.60 23 40 ô70 540 1270 \ M¡n¡rum 13 20 13 80 Cl ìÈ\

Ò

T6PHL T6PHR T6M1O .o T6M2 T6[41 T6M6 T6t!!9 T1 IFLCA T1 IFRCR T1 IFRCA 35 36 34 35 36 35 Valid 39 90 12.15 12 s6 16 67 01 26 25 27 10 87 650 t0 56 21 27 9s 12 20 12 60 16 70 '10 21 20 19 10 26 05 10 70 630 70 Med¡an 11 80 16 80 fl.704 25 50 25 204 12 20 1o 504 610 I 8oã 21 60 Modê 126 b 166 244 227 1 13 104 'I 50 96 146 136 \- std Oeviat¡on '14 23 60 980 10 60 50 510 720 t6 30 15 20 21 00 M¡n¡dm 760

TlOMg fIOPHL T6IFLCA T6IFRCA T10t\.,12 T1OMl Tt0M6 T6lr1 1 T6SPL T6TPW 40 38 34 13 N Valid 36 34 66 1572 12 67 23 83 22 24 31 25 17.86 18 70 65 50 13 35 3t 30 35 80 15 90 13 45 12 60 23 95 22 50 17 45 't7 60 66 00 M€d¡an 14 90 23 204 23 60 28 80 35 8oa 1o soa 55 5oa p 4oa 12.204 Mode f 610 152 324 337 143 544 444 1.76 128 146 Std Dev¡at¡on 147 17 80 2,1 50 25 20 12 60 '10 55 50 930 18 00 Min¡rum 13 80 50

o\OJ o\ ¡ S 110M10 T1 OSPL ll0tFLoA TlOIFRCA N Valid 38 40 11 24 36 36 15 84 16 44 l8 60 30 41 63 04 '13 06 1294 I 25 45 32 92 Median 15.95 16 30 18 50 29.70 63 80 't270 o 13 05 2A 20 25 60 33 30 Mode 16 1oâ 16 30 18 00 26 60 65 20 13 00 12 50 goa o 28 1oa 23 27 60e Sld Dev¡at¡on 122 158 180 374 486 t81 196 161 216 2AA G Minimm 13.40 13 20 14 10 26 40 48 00 ì 860 910 22 70 19 80 27 60 õ 16 40 ì o .o L1M9 L1 PHL LI PHR LlM1O LlMl1 LlSPL LlTPW L1 IFLCR Ll IFLCA L1 IFBCR G N 36 34 35 8 s4 31 Mean 40 99 16 36 't646 18.25 24 37 32 29 75 07 858 13 06 892 Mêdian 41 00 16 40 16 45 18 75 24 20 32 10 76 60 855 't3 00 890 L¡ode 33 80 15 20å 16loa 18 8oa 21 90 22 10ã .t4 76 60 890 1s 20 a 604 u Sld Devialion 338 131 163 1,71 202 611 tt 45 'I 30 232 129 Minill1h 32 30 f 4 t0 12 00 14 50 20 00 22 10 40 20 530 800 630

L5M2 Lst\,t1 L5M6 L5M9 L5PHL LsMlO L5M1 1 LsSPL N Valid 34 36 32 38 38 37 35 38 11 Mean 12 99 24.O9 28 92 34 53 17 73 13 92 't4 55 17 73 26 27 29 93 Med¡an 12 95 21 30 29 10 34 90 17.70 13 80 't4 50 17 70 26 65 30 60 l\.lode 11 10 22 60 23 10 33 204 ,t3 60 13 80 14 80 16 20à 23 20 24 60 Sld Dev¡alion 199 193 231 3.10 'I 510 62 185 229 304 389 Minirum 830 20 10 20 50 2A 20 3470 10 40 10 90 12 60 20 20 24-60

o) o\ ! HCMT FHBM18 FM¡¡16 FMMT L5IFRCR L5IFRCA 39 L5IFLCR 36 40 - 36 2A 38 37 67 2A 49 39 18 32 39 325 50 N Val¡d 9.79 37.30 653 10 09 6.35 49 10 91 45 3275 325 00 68 00 Meen 610 10.00 38 05 6 ,10 10 00 49 10 I 93 80 32s 00 6s 00 Median 33 3oa 29 704 6 ooa s.eo" \J 5.304 8.20å 318 79 80 225 14 53 555 Mode 1.71 258 227 .96 40 80 760 .97 286 00 56 00 a Std Dev¡at¡on 30 60 28 60 550 470 6.70 79 80 450 Min¡rum

ì o BIWM2 .o FLMl FCMS 24 36 39 Valid

tr

OJ æ ìt S Females - Modern

C3SPL c3M2 c3M6 c3t\49 C3PHR c3Mt0 c3t\411 25 26 24 26 I N 30 26 24 '16 09 622 625 15 44 23 86 03 U Mean 173 1277 12 52 14 72 18 60 14 80 17 80 640 635 15 50 23 90 14 o Med¡an 150 12 95 12 45 17 80 s go' 4AO tg zo' zz ao^ tz so" Mode 100 13 30 rz oo" 14 10 93 98 148 158 405 ì Std Dev¡at¡on 83 121 141 237 460 4AO 13 10 20 90 12 30 Minimm to0 10 30 10 40 11 a0 13 70

o .o

cTMl c7M6 c7t\49 C?PHL C3TPW CsIFLCA CAIFRCA c7M2 26 25 26 26 N Valid 17 27 12 76 15 97 24 37 650 È 51 85 667 a27 661 813 13 63 815 13 75 12 90 16 30 25 10 645 Med¡an 53 00 685 820 680 go' 790 13 40- 12 20 16 80 zs zou s sou Mode 53 30 s s zo' 730 97 '1 132 141 140 150 186 Sid Deviat¡on 08 125 105 590 10 70 10 30 l3 40 19 80 480 Minirum 42 20 430 580 430

CTIFRCR CTIFRCA T1t\,,t2 c7M1 0 CTSPL cTfPw cTtFLcR CTIFLCA 24 2A 27 26 27 14 6 26 26 1574 26 11 52 88 661 996 676 981 Mean 662 14 51 25 74 15 65 26 35 61 60 660 10 15 690 ss5 Median 650 14 60 25 60 2o 90" 22 zo' o so" 930 720 10 40 14go' Mode a zo' 14 60 z¿ tou 21 79 85 131 84 117 138 Sld Devìation 108 134 178 286 440 610 520 700 13 10 M¡n¡dm 480 11 40 22 20 20 90 22 20

CÐ \oo\ S T1M1 T1M6 T1M9 T1 PHL T1 PHR T1 MlO T1t\.,t11 TlSPL TlTPW TlIFLCR Þ N Val¡d 27 28 2A 2A 2A 13 2A

Mean 1,1 .16 16.01 26 10 a37 427 15 33 22 21 29 13 72 53 661

I 'l,t Med¡an 30 16 30 26 10 860 845 15 30 29 80 73 50 650 o '!6 Mod6 13 40 soa 21iOa 6 7oa 6 7oa 14 204 20 204 23 604 70 90ã 650 o Std Dev¡at¡on 140 180 204 127 108 118 168 3_02 390 'I 0l \ Min¡mm 1160 12 20 21 20 620 640 12 E0 18 90 23 60 63 50 440 ì o -o T1 I FLCA T1 IFBCR T1 IFRCA T6M2 T6M1 T6PHR T6M1O

N Val¡d 27 28 27 27 2A 27 27 26 \ M€an 10 34 6.f2 10 42 19.83 17 73 23 57 21 60 10 54 10 80 16 16 Median 10 60 675 't0 50 19 70 17 80 23 50 24 35 t0 30 f0 70 16 25 Ë Mod€ '11 20 5 goa 10 20_ '18 90 16 80 24 20 23 ooa lo 20 I 60å 15 30 b \- Std Dev¡ation 134 92 1.54 140 117 231 196 125 1.20 102 M¡nirum 810 450 630 17.50 '15 80 20 60 20 20 860 860 '13 90

T6M1 1 T6SPL T6TPW T6IFLCA T6IFBCA T1OM2 T1OM1 T10t\¡6 T1OMg TlOPHL

N Val¡d 11 22 24 25 2A 2A 26 2A

M€an 16 88 17 01 60 93 12 42 11 47 22 05 21,45 27.2A 30 98 14 27

Med¡an 17 10 't630 61 25 12 3s 12 00 21 95 21 15 26 80 30 70 14 25

Mode 15 l0a 7 80â 62 60 10 50- I 50â 1g 60_ 20 60 2s ßa 30 00ã P 4oa Sld. Deviat¡on 175 564 423 197 169 't 45 175 247 235 170

Mihimm 13 80 7.80 54 20 7.70 830 19 60 16.40 2290 26 10 11 20 7n ln

(¡) N O 1,1M6 TlOIFBCA Lf L1M1 T1 0M1 1 TlOSPL TlOPHR T1 T1 25 2A 13 * 27 24 97 2A 17 '12 62 26 44 N 58 08 66 12 't7 85 2ô 05 14 34 16 44 10 27 AO 12 60 26 80 2s 26 90 s7 95 12 A5 60 17 f5 I 14 20 16 22 20' 29 40 12 104 26 80 Med¡an 49 6oa p 20à 16 704 20 20À \J 11 90ã 15 8oa 1 8,1 261 1 15 204 413 112 168 130 171 21 70 24 00 o Sld Devial¡on 00 10 70 22 40 20 20 49 60 10 13-50 14 80 ì M¡niltln 11 90 e) ì o IiFLCR LlIFLCA L1 LtM11 LlSPL L1 _o L1M9 PHL LlPHR 27 13 t5 2A 28 13 56 948 Valid 28 68 30 9.12 23 18 29 24 14 63 1A 42 35 92 14 36 915 13 90 ts 23 45 30 20 7020 65 '18 35 14 20 14 14 80 920 3s 70 57 6oa 6 7oa Med¡an 23 90 28 50ã 12 AOa 17 80 34 goa ß 204 137 145 130 Modê 177 399 6.53 153 155 b 280 '1.36 10 30 720 Std Deviation 20 00 57 60 670 \- '11 20 00 30 60 12.1 0 10 Minirum

L5t\,t11 L5PHR LsM1O L5M6 L5M9 L5M2 13 26 26 21 23 27 26 52 2674 Valid 13.29 17 71 26 N 42 57 12 65 26 12 30 38 '13 43 23 60 26 00 26 80 95 1275 17 30 30 50 43 10 f2 23 60 2ø 10 23 604 13 50 15 90ã 24 204 Med¡an 10 zoa 13 ooa 12 60á 26 SOa 30 50â 1t 80ã 22 404 293 406 223 209 190 208 268 322 135 179 21 90 19 50 Std Deviation 750 880 14 70 2370 25 60 36 90 11 10 1 9.80 Minirum

ol\ ì

S 151 L5IFLCR L5IFLCA LsIFBCR N L5IFRCA FMt\4f ã 15 6 FMt!,t7 27 HCMT FHBIú1 8 Mean 27 27 a4 4A 695 21 29 1t 31 713 30 29 Msdian 11 2A 35 84 I a2 40 690 31 05 304 28 11 20 720 58 67 43 85 i,lode 11 20 36 20 o a2 10 30 90 305 00 580 11 00 58 00 43 80 Sld. 530 't050 D€vlation 38 30 28 30ã lo 56 1,14 308 00 o 164 59 00 43 6oa M¡n¡mh 128 1s8 62 30 349 2.7ã 530 730 21 16 501 s Mâyìñ,m 440 870 17 I 10Õ 10 30 l0 27 00 890 266 00 50 00 ñ -l¿ 80 510 37 20 \

o

.o FLMl FCMs BIWM2 N Valid 26 R 29 25 Mean 426.54 82 66 242 16 Med¡an 429 50 83 00 280 00 Mode 436 00 85 00 270 00 Sld D€vlat¡on ¿ 25 27 609 Minim 15.69 373 00 î 71@ 248 00

a Muftiple mdes exist. The s@¡lest val@ ¡s slþM

(¡)\ NJ 4. Sexual dimorphism of all variables in modern samples (means of modern samples, percentage-diffelence, t-values)

males Mean females males > females ($ Variable Mean

14.6 12.8 1 1.03 s.31 C3 dorsal vertebral body height dorsal 't0 't4 12.s 62 5.57 c3 ventral veñebral body heighr 1 16.2 14.7 744 4.01 C3 vertebral body sagittal diameter :: r 8.1 3.97 2.14 Cs veftebÊl body transverse d¡ametet 19.3 13.68 4.33 C3 left ped¡cle height 7.3 6.3 12.08 3.90 Cg r¡ghr pedicle height l5 4 2.35 1.28 C3 spinal canal sagìttal diameter 23.9 420 166 C3 spinal canal transverse diameter 245 '16.9 16.0 17.7 4 0.60 Cg spinous process lengll 56.1 51.9 8.81 3.51 C3 transverse Process width 1.15 061 C3 left cranial inlervertebral foramen width 68 -0.90 018 ' C3 left caudal interuelebral foramen widlh 8.2 6.6 -1.76 0.48 C3 right cranial inlervertebral foramen width 65 8l -2.48 0.21 ' C3 righl caudal intervertebral foramen w¡dth 82 13.6 869 4.48 C7 dorsal vertebral body height 15.2 12.8 729 2.65 c7 ventrcl veftebral body height 13.7 17.7 16.0 914 4.17 C7 vertebral body sagitlal diametel 26.6 244 6.33 4.25 C7 venebral body transverse diameter 6.5 8.73 4.29 C7 left pedicle he¡ght 7.5 6.6 ðJU 3.57 C7 right ped¡cle height 3.80 1.65 15. 1 14.5 C7 sagitlal diameter spinâl canal 25.7 3.09 0.93 C7 lransverse diameter spinal canal 26.1 31.5 26.1 13.04 4.72 C7 spinous Process lenqlh 662 529 17.64 r.29 C7 transverse process width 6.6 -3.30 135. C7 lell cranial interuenebralforamen width 6.3 't0.1 10.0 1.87 026 C7 left caudal inlervertebral loram en width 6.6 6.8 -0.90 099. C7 right cranial intervenebral foramen width 9.S 3.18 0.96 C7 right caudal ¡nlerveneblal loramen widlh 10.1 15.7 8.97 4.59 TH1 dorsal vertebral body heiqht 17.3 16.0 14.5 9.11 4.67 TH1 ventral ve¡tebral body he¡ght 16.0 8.75 3.90 THI sagittal diameter vertebral body 178 26 I 7.96 4.&r TH1 lransverse d¡amelel vertebral body 28.9 9.3 8.4 1 0.20 3.06 THl left Ped¡cle heighr 9.1 8.3 922 3.00 THl right Ped¡cle height 't5 15.3 3.27 1.52 spinal canal sagiltal diam eter I TH1 516 2.50 THl spinal canal transveße diamelet 29.1 13 01 3.66 length TH1 spinous Process 5.72 79 1 72.5 907 TH1 l¡ansverse Process widlh 6.6 042 0.20 TH'l lelt cranial interuenebral foramen width 't.53 10.9 10.3 0.94 THI lett caudal inteFr'ertebrel foramen width 6.5 6.7 -0.70 0,97. THI right cranial interyenebral loramen width 10,6 10 4 0.76 0.38 TH1 r¡ght caudal inleruerlebral foramen width 19.8 844 3.35 TH6 dorsal vedebral body height 21 0 '19.0 17.7 779 3.56 TH6 ventral vertebral body he¡qhl ZJþ 10 59 4.46 TH6 sag¡ttal diameler vertebral body zÞ.J 27.9 246 10.98 6.34 TH6 lransverse diameter vertebral body 10 5 13 22 5.33 TH6 left pedicle height 126 10 s 13.86 6,19 TH6 ¡ight Ped¡cle he¡ghr 167 16.2 2.45 1.76 TH6 spinal canal sagitlal diam eter 17.7 169 4.33 1 .89 TH6 spinal canal transverse dlameter 18.7 170 17.81 078 TH6 spinous plocess length 65.5 60.9 8.28 3.74 TH6 transverse width Process -0 1.85 I ó.J 12.4 8r TH6 lefl caudal intervenebral foramen width 1 1.5 0.71 3.01 TH6 r¡ght caudal inleruertebral foramen w¡dth 12.7 22.1 820 5.01 TH10 dotsâl vertebral body height 23.8 22.2 21.4 5.96 1.92 TH10 ventral vertebral body height 31 3 12.42 5.31 TH10 vertebral body sag¡ttal d¡ameter '1.07 34.7 31.0 1 5.39 TH10 venebral body transvelse diameter 14.3 10.09 TH10 left ped¡cle heiqht 157 l4 3 9.32 4.01 THl0 r¡ght pedicle height 15I 16 4 16.4 2.91 0,01 TH10 spinal canal sagiltal diameler 18 6 17.9 5.67 176 TH l 0 spinal canal lransverse diam eter 26 1 9.68 283 TH10 sp¡nous ptocess length 304 63.0 58.1 9.33 3.59 THlO transverse Process w¡dth 't3.1 12.7 087 Ll0 TH10left caudal interyertebral loramen width 12.9 12.6 132 083 TH10 rlght caudal intervertebral loramen widlh tAL 6.08 3.16 Ll dorsal vedebral body heighr 279 25.5 25.0 4.03 0.94 L1 ventral venebral bodY height 28.2 12.85 6,64 Ll venebral body sagittal d¡ameler 32.9 41.0 35.9 1 1.84 6,70 L1 vertebral body transvelse diameler 14.4 8.75 5.89 Ll left pedicle height 16.4 '16.5 14.6 9.22 4.68 Ll r¡qht Pedicle he¡ght - 18.4 0.42 Ll spinal canal sagittal diameter 182 24.4 232 4.96 2.55 Ll sp¡nal canal tÊnsveße d¡ameler 323 29.2 12.32 1.34 L1 spinous process length

F. J. RühIi - Osteomelric Variatiott of tlrc Htunan Spine J/J Ll lransveße process widlh 68.3 1 1.49 2.11 Ll lelt cran¡äl interveñebral loramen width 8.6 9.1 -7.27 1.60 ' -3.43 L1 l€ft caudal intervertebral foramen width 13.1 13.6 1.05 ' -5.92 L1 right cranial ¡nteryertebral loramen width 8.9 9.5 168t -2.58 L1 r¡ght caudal interuertebral foramen width 13.0 13.4 1.04 ' '16 L5 dorsal vert€bral body hêighl 24.1 23.6 4 1,04 L5 ventral verlebral body heighl 289 28.1 5.ô l 1.43 L5 vertebral body sagittal diarcter 34.5 30.4 754 5.,1O L5 vcrlcbral body llansvcrac diamtcr 47.7 42.6 7.75 5.07 L5 lelt pedicle height 13.9 12.7 s56 2.54 L5 righl pedícle heighl 14.5 7.00 t Êl L5 spinal canal sag¡tlal diameler 17.7 17.7 o.o7 005 L5 spinal canal transverse diqm€ter 26.3 0.87 0.34' L5 sp¡nous proces1us lenglh 29.9 7.79 2.05 LS lønsveße prccess width 91.5 84.5 a47 L5 left cran¡al interuertebral loramen wìdth 6.5 1.56 ' Ls left caudal ¡nleryedebral loramen widlh 10.1 -5 40 Ls tight cÊn¡al inleNedebral foramen widlh 6.3 -6 57 2A9' L5 r¡ghl caudal intcrycrtcbral lorarcn widlh 9.S 1 1.3 -6 60 3.61 ' loramen magnum sagittal diameter 37.3 35.8 397 1.65 loramen magnum transverse diameter 32.4 31 0 6.35 '1.86 huærus lcngth 325.5 304.3 4.28 4.67 huæfuÊ circunlerence 67.3 58.7 12.A4 6.89 lcmral hèad w¡dth 494 43.9 1 1.70 7,29 lcm? lcnglh 453 4 426.5 4.21 tcm? c¡rcurlcrcrcc e9.4 82.7 10.90 4.35 bi-¡liac width 289.8 282.2 4.00 1.68

t: b¡gger mean value in f€målês than mal€s Ital/c: significant (p<0 05) before Bonferroni's co116ct¡on Bold: significant (p<0 05) after Bonferoni's correction

F. J. Ri¡hli - Osteometric Variation of the Human Spine 374 5. Side differences of variables in modern samples (means, t-values)

Variable Males Females Mean (mm) t Mean (m m)t

C3 left pedicle height 7.3 6.2 C3 right pedicle height 7.3 0.27 6.3 0.12 C3 left cranial intervertebral foramen width 6.8 6.7 C3 right cranial intervertebralforamen width 6.5 1.58 6.6 0.20 C3 left caudal intervedebralforamen width 8.2 8.3 C3 right caudal intervertebral foramen width 8.2 0.04 8.1 0.40 C7 left pedicle height 7.5 6.5 C7 right pedicle height 7.5 0.01 6.6 0.43 C7 leÍl cranial intervertebral foramen width 6.3 6.6 C7 right cranial intervertebral foramen width 6.6 1.28 6.8 0.64 '10.1 C7 left caudal interveftebral foramen width 10.0 C7 right caudal intervefiebralforamen width 10.1 0.1 5 9.8 0.45 TH1 left pedicle height 9.3 8.4 TH1 righ pedicle height 9.1 0.68 8.3 0.34 TH1 left cranial intervertebralforamen width 6.6 6.6 TH1 right cranial intervertebral foramen width 6.5 0.28 6.7 0.45 '10.9 TH1 left caudal intervertebral foramen width 10.3 TH1 right caudal intervertebral foramen width 10.6 0.93 10.4 0.22 TH6 left pedicle height 12.2 10.5 TH6 right pedicle height 12.6 1.60 10.8 0.77 TH6 left caudal intervertebral foramen width 13.3 12.4 '1 TH6 right caudal intervertebral foramen width 12.7 1.79 1.5 1.85 TH10 left pedicle heighl 15.7 14.3 TH10 right pedicle height 15.8 0.39 14.3 0.17 TH1 0 left caudal intervertebral foramen width 13.1 12.7 THlO right caudal intervedebral foramen width 12.9 0.28 12.6 0.15 L1 left pedicle height 16.4 14.4 L1 right pedicle height 16.5 o.27 14.6 0.70 L1 left cranial intervedebral foramen widlh 8.6 9.1 L1 right cranial intervertebral foramen width 8.9 1.08 9.5 1.02 't L1 left caudal intervertebral foramen width 3.1 13.6 L1 right caudal intervertebral foramen width 13.0 o.12 13.4 0.33 L5 left pedicle height 13.9 12.7 L5 right pedicle height 14.5 1.58 13.3 1.08 L5 left cranial intervertebral foramen width 6.5 6.9 L5 right cranial intervertebral foramen width 6.3 0.85 7.1 0.55 L5 left caudal intervertebral foramen width 10.1 11.3 L5 right caudal intervertebral foramen width 9.8 0.64 11.3 0.07

375 F. J. RühI¡ - Osteometric Variation of the Humnn Spine 6. Correlations of variables with individual age in modern samples

Modern - males

AGE ACE Tllt{z 22t c3w Pemon Corelstion 341+ N 40 N l8 TIMI Peùson Codelålioû 138 c3Ml Pffion corelat¡on 285 N 38 N 38 TI M6 Peilson Cor€låtion .708r. c3M6 Pffion Corelå(ion 682++ N 34 N 36 TI M9 Peesoû Comlâ(ion 435r+ c3 M9 Peffion Corelat¡on 623.+ N N 37 TIPHL Peùson Corelation 393+ C3PHL Pemon Corelâtion 500.. N 40 N t5 TIPHR Pemon Corelât¡on 438r+ C3PR Pe{sonCoÍeld¡on .539++ N t9 N 33 TIMIO Peüson Coreldion 206 c3Ml0 Pemon Corelålion - t21 N 38 N l4 TIMI I Peffion Corelåtion 302 c3Ml I Peffion Corelãt¡on 345. N 40 N 38 TISPL Peffion CorelåLion 093 C3SPL Pffion Corelåtion 584 N 20 N ll TITPW Pffion Corelåtion 527 rt C]TPW Pwson ftrelat¡on 442+ N 35 N 22 TI IFI¡R Peüson Corel¡tion - 08l c3IFrcR Pemn cor€låtion -2t9 N 38 N 37 TIIFIIA P€ùson CoEelalior I t8 cslFt-cA PemonCoreldion -,o37 39 N t5 TI IFRCR Peùson CoÍelålion -o5l C3IFRCR Pemof, Coreldion - 122 N 35 N 34 TI IFRCA Psson corelat¡on 163 C]IFRCA P€ùson CoFelation ol I N 39 N 35 T6M2 Pøson Corel¿t¡on 288 clkf2 Pemof, corelåt¡on 221 N 36 N 38 T6MI Pffion Coreldion 133 clMl P€mon Coûelålion 242 N 35 N T6M6 Pffion Coreldion 483+. c7ïre Pemon Core¡d¡on 627.. N 74 N 36 T6M9 Peffion Coreldion 253 glw P€ffion Coreldion .451+r N 3ó N 16 T6PHL Pemon conelåtion 424+ C?PHL Peeson CoFelåt¡on t20 N 35 N 36 T6PHR P€mon Greldion 508++ OPR Ptroû Conelil¡on 231 N N l8 T6MIO Pøson Coreldion 189 c7Ml0 Person Coreldion 004 N 34 N l8 T6MII Peùson CotrelâLion - 061 c¡Mu Pøson Coreldion - o21 N 36 N 3E TóSPL Peeson CoÍelaLion -.404 CTSPL PeñoûCo[€laLion 150 N l3 N 23 T6PW Peüson Corelution 250 c7'IPW Pffion Corelåt¡on a3t+ N 27 N 1 T6IFI¡A Peilson Corelation t60 CTIFI¡R Pemon Corelat¡on - 25t N 32 N 35 T6IFRCA Peilson Coreldion 108 cTlFlSA Petrson Corelãtion 158 N 32 N 38 TIOM2 Pemon Coreldion z16 g¡IFRCR Pëùson Corelation - 262 N 38 N 15 TIOMI PeffionCorela(ion 056 C?IFRCA PeffioncoÍeldion 136

F. J. RühIi - Osteometric Variation of the Human Spine 376

- AGE ACE

TIOM6 Pøson Corelåtion 31 tl TJPHL Pffion Corel¡tion 250

N 3l N

TlOM9 Peùson CorelåLion 555r1 I.5PHR PetrsonCorel{ion 325r

N 39 N 31

TIOPHL Peüson Corelation 60 TJMIO Pcmon corelÀt¡on -083

N 44 N 35 220 TIOPHR Peùson CoÍelalion tJMT I Pè6on Corelålion t62

N 3E N 38 Tl0Ml0 Pffion Corelålion 267 TJSPL Pemon Coreldion 45f

N 38 N

Tl0Ml I Pemon Coffeldion ot1 TJTP}V Pemon Corelation 24A

N 40 N 8 318 Pemon Corelåtion 446r* TTOSPL P€ffion Corelâ{ion L5IFrcR

N ll 38 088 TIOTPW P€ilson CorelaLion 288 LíIFrcA P€don Corelation

N 24 N 37

TIOIFT'A Pemon Õrelåt¡on 0ll IJIFRCR PúsonCor€lalion 262

N 38 N 35

TIOIFRCA Peffion Corelal¡on - 013 LJIFRCA Peffion ftFelat¡on - 149

N 31 N 36 Lrw Pemon Con€lalion s1 Pefion Corelation 259 N 36 N 28 LIMt Pmon Corelâtion 305 FMMT Peffion Cocld¡on -070

N 36 N 28

LIM6 Peùson Corelâtion .189 HLMI Petrson Cofi¿låtion 227 N 33 N 36

LIM9 Peüson Corel¡t¡on ,435++ HcNft Pffion Corel¡tion 524++

N l1 g

LIPHL Peffion Corelat¡on 340. N 15

LIPHR Peffion &relation 470.1

N 36

LIMIO P6on Coftldion 087 AGE

N 34 FHMl 8 Pearson corelal¡on 271 LIMI I PøonCorelilion 090 N 39 - N 35 FLMl Fearson Coríelalion 121 36 LISPL Pø5on Corelation 396 N

N FCMs Pearson Corelalion 421"

- LTTPW PeffionCoreldion 070 N

N l6 BIWM2 Pea6on Corelallon ,307

LlIFI¡R P€mon ftreldion - 043 N 34 ' Corelåt¡on is siFilicilt åt ú€ 0 05 lev€l (2{ailed) +r the0 lev€l LI IFIfâ Pemon coñelåtion o74 Corelålion ¡s significil! a! 0l (?-tailÈÐ N 34

LI IFRCR Peffion coFelation t20

N 12

LIIFRCÀ Pemon coreldion 28

N 34 IJMZ Pemon Cofieldion 2M

N l6

I.JMI Petrson corelå(ion l8l J1

L5Mó PeffionCoF€lttion 152t.

N 32

fJM9 Pemon ftnelâtion 6l6rr

F. J. Rühli - Osteometric Variation of the Human Spíne ó// Modern - females

AGE AGE .148 T1M2 Pearson Corelalion c3M2 Pearson Corelal¡on - 064 N N 26 Peaßon CÆrelalion -.236 c3M1 Peårson Corelalion - 287 T1M1 N N Pearson Corêlalion 013 c3M6 Pearson Corelalion 293 Tr ¡.,t6 N 27 N 25 Peareon Corelal¡on 092 c3M9 Peaason Corelalion 397' T1 M9 N 2g N 25 Pearson Corelal¡on 137 CsPHL Peárson Corelat¡on 165 fIPHL N 28 N TlTPHB Peaßon Corslalion 209 C3PHB Pearson Corelalion 171 N 28 N 26 PÉarson Corelal¡on 299 caMlo P6aßon CÆrelal¡on 074 flMt0 N 28 N 24 Pearson Corelal¡on 2't4 c3M1 I Pearson Correla(ion 167 TlM1I N N 26 Pearson Cotelalion - 383 CsSPL Pea¡son Corelal¡on 645 T1 SPL N 13 N s Pearcon Co16lal¡on .o77 æTPW Pea6on Corelalion TlTPW N N 17 Pearson Corelalion 056 CsIFLCR Peaßon Corelalion T1 IFLCB N N 26 TlIFLCA Peaßon Cotelallon 194 CAI FLCA Pea6on Correlalion - 046 N N 27 TIIFBCR Peaßon Corelalion o27 CsIFRCR PeaÉon Corelalion - 008 N 2A N 26 TI IFRCA Peaßon Corslalion 306 CsIFRCA Peaßon Corelal¡on 017 N 27 N T6M2 Peaßon C4relat¡on 037 c7M2 Peaßon CoÍ€lat¡on 106 N 26 N 26 P€a60n Corslat¡on - 036 C7MÍ Pearson Corelal¡on - 206 T6M1 N N 25 Peaason Corelalion 176 c7M6 P€arson CoFelalion 209 T6M6 N 27 N 26 T6M9 Pearson Corelãlìon - 234 c7M9 Pearson CÆrelation 088 N N f6PHL P6arson corelalion - 210 CTPHL Pearson Correlalion 132 N 27 N 26 f6PHB P6aßon Cor€lal¡on .065 CTPHR Pêaßon CÆtelalion 3S9' N 27 N 27 T6MIO Pearson Co(elalion c7M10 Pearson Corelation 322 N 26 N 26

l6M1 1 Pêarson Co(elalion c7M1 1 Psaßon Correlat¡on - o44 N 27 N Pearson Corelalion 30s CTSPL Pearson Coralalion ..397 T6SPL N 11 N 14 T6TPW Pearson CÆ(€lalion CTTPW Peaßon Corelalion . 575 N 22 N 6 fôIFLCA Pearson Corelalìon 108 CTIFLCR Pearson Corelalìon - 224 N 24 N 26 fSIFBCA Pêarson CÆrelalion 't 45 CTIFLCA Pearson Corr€lal¡on 299 N N 26 TlOM2 Pea60n Correlal¡on - 230 CTIFBCR Pearson Corelation 105 N 2g N T10Ml Pearson Corelalion - 244 CTIFRCA Peaßon Corelal¡on 243

378 F. J. Rühli - Osteometric Variation of the Human Spíne AGE AGE Pearson Correlalion 224 f'r0M6 Pea6on - o55 L5PHL 26 N '185 - L5PHR Poarcon Corelat¡on T1OM9 Peã6on Corelal¡on 066 N N ! L5MlO Pearson Colelatlon .086 TlOPHL Peaßon Corelation 2so N 25 N 28 129 129 LsM1 1 Poarson coarÊlalion T1 OPHR Pearson Cotelalion N 27 N 27 Corelalion .,043 ft0M10 P€arson CÆtelalion 288 L5SPL Pearson N N 27 Cor€lat¡on . 153 Tt0Mll Pêarson Conelal¡on 066 LsTPW Pearson N l5 N 28 -,o74 L5IFLCB Peaßon Correlålion 168 TIOSPL Pearson Corelal¡on N N 13 214 186 LsIFLCA Pearson Col6lalion TI OTPW Pearson Corelalion N 27 N 22 Peerson Correlal¡on .037 TIOIFLCA PeaÉon Corelalìon 117 L5IFFICFI N 27 N 2A Pearson Colelalion 135 TlOIFRCA Pêarson Corslalion 187 LsIFRCA N 27 N 090 - FMMl6 Pêarson Corelalion L1M2 Peaßon Corelal¡on 086 N 21 N 27 .138 - 338 FMMT P€arson Corelat¡on L1M'I Peã6on Corehl¡on N N -.343 HLMl Pêarson Correlalion L1M6 Pearson Cot€lalion 213 N 2S N 25 -.s46 - 096 HCMT Pearson CÆrelalion Ll M9 Peaßon cotelalion N 30 N 2ø - 199 FHMl 8 Pearson Col€lal¡on LlPHL Peaßon Corelalion ,240 N 22ê L1 PHB P€aßon Corelalion

N 2g

P€aßon Correlal¡on o31 LlMlO AGE 2E N -_474. FLMI Ptron Corelåtion 11Ml1 Pea6on Corelallon 331 N 26 N 28 - 359 FCM8 Pemon cotreldion LlSPL Pearson Corelalion 106 N 29 N t4t BtwM2 Pemon Corelâtion LlTPW PsarÊon Côr€lat¡on - 060

N l5 (2ldl€d) ' Coreldion is signifimt al theO 05level L1 IFLCR Peatson corrêlalìon

N 2A - 050 L1 IFLCA PeaFon Correlalion N 261 L1 IFRCR Pearson Corelalion

N 27

LIiFRCA Pearson corelal¡on 130 N 27 - 204 L5M2 Pearson Corelallon

N -,479' L5MI P€arson Colelâtion

N 071 L5M6 Pearson Cor€lâlion

N 23

L5M9 Pearson Corolal¡on .s78

379 F. J. Rühti - Osteometric Variation of the Human Spine 7. Correlations of variables with major age groups in non-modern samples

Non-modern - males

AGEGROUP AGEGROUP Colelal¡on ,136 c3M2 Pea6on CÆrelalion 162 TIM2 Pearson

N a7 N Pearson Corelal¡on 201 c3M1 Pea.son Co(elal¡on 182 TlMI 93 N 06 N CÆt€lation 468" c3M6 Pearson Corelal¡on T1M6 Pea6on 90 N 83 N Colelallon 21ø' æM9 Pearson Corelalion - o70 flM9 Pearson 93 N 80 N 217', C3PHL P€arson Corelal¡on 240' T1 PHL Pearson Corelat¡on 88 N a2 N Cotelalion CsPHB Pearson Corelalion ,262' fI PHR P€arson 89 N N Córrelation 010 c3M10 PeaEon CÆrelal¡on ..062 TIMIO PeaÍson 84 N ?3 N 03t - os1 TTMl 1 Pearson Corelal¡on c3M1 1 Pearson corelalion 85 N 8l N .028 Cotrelat¡on 204 C3SPL Pearson Cotelatlon TlSPL Pêaßon

N N Corelalion 269' CSTPW Pearson C¡rrelat¡on -o22 fITPW Pearson

N 50 N 6f f78 CAIFLCB Pearson Corelalion - t25 fl IFLCR P€arson Corlelat¡on 86 N 8l N - 055 C3IFLCA Pearson Correlal¡on - o17 T1 IFLCA PeaÞon corelal¡on 85 N 81 N Coilslal¡on - 108 CSIFRCB Pearson Corelalion - 15t T1 IFRCR Pearsoo a2 N 85 N Corelalion - 120 CAIFRCA Pea6on Coíelation - 070 T1 IFRCA Pearson a2 N 84 N Correlallon .400" c7M2 Pearson Corelãl¡on - 019 T6M2 Peaßon g8 N N Co(elat¡on 180 c7M1 Psarson Corelal¡on 035 T6M1 Pêaßon

N 90 N Cotelat¡on 469" c7M6 PeaFon Corélal¡on .594- T6M6 Pearson 82 N N Co(elal¡on .338" c7M9 Pearson Core¡alion 356- T6M9 Pearson

N N Corelal¡on 219' CTPHL Peãßon Corelallon 098 T6PHL Pearson 83 N 85 N Pearson Corehl¡on 256' CTPHR Peaßon Corelatlon 069 T6PHR 85 N 85 N . Pearson Correlallon 003 c7M1 0 P€arson Corelat¡on 053 T6MlO 74 N N 080 T6M1 1 Pearson Corelalion c7M1 1 Peaßon CÆrelal¡on 174 84 N 90 N Pearson Corelat¡on - !50 CTSPL Pearson Correlal¡on 122 TsSPL

N N 349' CTfPW P€arson corelallon o52 T6TPW Pearson Cor€lation 43 N 26 N 032 CTIFLCR Pearson Corelat¡on - 112 T6IFLCA Pearson Colrelalion 6S N 86 N Colêlal¡on -.17 4 CTIFLCA Pearsoñ Corelalion -,120 T6IFRCA P6aEon a7 N 83 N Correlalion .215' CTIFRCR Pearson Correlal¡on -,203 TlOM2 Pearson 97 N N Pêarson Corr€lal¡on '167 CTIFRCA Peárson CÆrelal¡on -11r T1OM1

380 F. J. RühIi - Osteometric Variatiott of the Human Spine AGEGROUP AGEGROUP

T1OM6 Pêarson Coilelation 468" LsPHL Pearson Correlallon 198'

N 93 N 101

TIOM9 Pêarson CÆrelat¡on 366" LsPHR Peaßon CoÍelation 221

N 96 N 98

TlOPHL Psarson Corelalion 185 L5M,IO Pearson Corelalion 133

N N 86

TlOPHR Pearson CÆt€lal¡on LsM1 1 Pearson Corelalion 170

N N 92

T10Mt0 Pearson CoÍelal¡on - 004 L5SPL Peaßon Corelalion -3lt'

N 89 N 44

TtoMft Poarson Corelalion .037 L5TPW Pea6on Coraelalion 051

N s3 N 40

T1 OSPL Pearson Cor€lãl¡on 258 L5IFLCR Pearson Co(elalion - 137

N 42 N 87

T1 OTPW Pearson Cotrelat¡on 231 L5IFLCA P€arson Correlalion

N 57 N B7

T' OIFLCA Pearson Corelal¡on 163 L5IFBCF Pearson Conelalion .018

N 86 N s3

TlOIFRCA Pêaßon CÆre¡alion -.087 LsIFBCA Pea6on Conelalion -,o23 a7 N a4

Ll M2 Peãßon Corelalion 118 FMM1 6 Pearson Corelalion 231

N 111 N 28

L1 M1 Psarson Corehl¡on 150 FMMT P6arson Corsht¡on 017

N '|05 N

L'IM6 Pearson CoÍelal¡on .319- HLMl Pearson Corelal¡on 292"

N 99 N 90

LîM9 Pearson Corelalion 299* HCMT Pea6on corelal¡on .219'

N 110 N 117 Peãrson CÆrelal¡on Ll PHL PeaEon Cotrelalion 191 FHBMl8 .376-

N 106

Ll PHR Psaßon Corelal¡on 140

N 107 LlMlO Pearson Corelal¡on 060 AGECROUP FLMI Peüson corelÂtioo 329+1 N N 94 L1Ml 1 Pêarson corelalion 206' FCMs Person Coældion 215++ N 102 N tû LlSPL Paarson Corelal¡on 103 Peùson Corelat¡on -165 N 46 BIWIV{2 LlIPW Pearson Corolâl¡on .293 Corelatlon is sign¡licant al lhe 0.01 level (2-lailed)' N 36 " ' Correlalion ls sign¡l¡cant al lhe 0 05 level (2la¡led)' LI IFLCB Peaßon Corelal¡on 079 g9 N

Ll IFLCA Pôaßon Corelal¡on - o12

N s3

L1 IFBCR Pêã6on Coarelalion - 05t

N 95

Ll IFRCA Pearson Corelal¡on 081

N 92

L5M2 Peaßon Corelalion 186

N 100

L5M1 P6arson Corelalion 240'

N 102

L5M6 Pêarcon CÁrrslal¡on

N 100

L5M9 Pearson Corelalion 277"

F. J. Rühli - Osteometric Variation of the Human Spine 381 Non-modern - females

AGEGROUP AGEGROUP 172 c3t/2 Pearson Co(elalion 079 T1 M2 N s8 N 'l02 Corelal¡on - 029 caNtl PeãEon Correlal¡on T1M1 Pearson N s6 N 101 Pearson Coil6lal¡on 354" c3M6 Pearson Cor6lalion 421" T1M6 92 N 9g N - Pèaßon Co(elal¡on 240' csM9 Pearcon Cote¡alion 026 T'M9 N N PeaFon Co16lal¡on 174 C3PHL PoaEon corelôllon 148 T.I PHL N s3 N 120 T1 PHR Pêatson Cor€lal¡on CAPHR Pearson CoÍelal¡on 292* N 98 N 98 .,213 Pearson Corelation .094 csMto Psarson Corelal¡on TlM,IO N N 77 P6aßon Co16lal¡on '179 csM11 Pearson Corelat¡on 035 T1M1'I N N s7 Pearson Corelat¡on -,217 C3SPL Pearson Corelal¡on 37 4' TISPL 40 N 36 N Peaßon Corelation ,o22 CATPW Peareon Corelal¡on .081 TlTPW N 64 N CÆrelal¡on -.1 56 C3IFLCR Peaßon Cotrelat¡on - 220' T1 IFLCR Pearcon 88 N 95 N - Pearson Corelalìon - 082 CAIFLCA Peaßon Cotrelal¡on 189 T1 IFLCA N N 96 PeaFon Corelalion - 226' CsIFRCR Peaßon Corelalion - 192 TlIFRCR N 90 N 97 Pearson Corslal¡on .,036 C3IFRCA Peaßon Cofi€lal¡on 103 T.I IFRCA N ø7 N Pearson Corelal¡on 071 c7M2 Pearson Corelal¡on .o75 T6M2 95 N 100 N - PÉaßon Corêlal¡on - 030 c7M1 Pearson Cotelal¡on o45 T6M1 95 N 101 N Pêarson CÆrelation 391 - c7M6 Pearson Corelal¡on 422" T6M6 N 93 N 99 Corelalion 153 c7M9 PeaFon Corelalion 198' T6M9 Peaßon N N 99 P€aßon Cotelal¡on 156 CTPHL Pearson Cotelalion .116 T6PHL N 91 N 97 Pearson Cotelal¡on î50 CTPHB Pearson Co(€lal¡on 113 T6PHB N N 101 Pearson CÆrelalìon - 051 c7M10 Pearson Corelatlon o2a T6M1 O N N 93 102 .069 T6M1 I Pearson Coûelalion c7M1 1 Peaßon CoÍelation N 91 N 95 - Pearson Corelat¡on - 239 CTSPL Peaßon Cotrelalion 199 T6SPL

N 55 N Pearson Corelal¡on 241 CTTPW P€aßon Corelallon 247 f6TPW N 65 N 27 .,331 Pearson Correlal¡on 010 CTIFLCR PâaFon Corelat¡on " T6IFLCA N 74 N 91 .,o72 Psarson Cot€lalion 015 CTIFLCA Pearson Co16lalion T6IFRCA N 72 N 88 Pearson Correlalion 060 CTIFRCB Pearson Coralalion - 232' TlOM2 105 N N Peârson Corelalion - 014 CTIFRCA Pearcon CÆrelalion -,228' TlOM1

382 F. J. Rühli - Osteometic Variation of the Human Spine AGEGROUP AGEGROUP 079 LsPHL P€arson Pêarson Corelal¡on 0s2 TlOM6 104 N N 104 - 034 L5PHR P6arson Coíelalion P€aason Co(€lation 159 T1OMg 106 N N 105 Corelalion - 019 't27 L5¡¡10 Peaßon Pearson Corelal¡on TlOPHL 92 N N 101 Pearson Corelalìon 117 .039 L5¡,,111 TlOPHR Peaßon Corelalion 101 N N 103 141 LSSPL Pearcon Colelalion Peatson Corelal¡on o52 T10M10 46 N N 95 008 ..036 LsTPW Peaßon Colelal¡on T10M11 Pearson Cotelalion 55 N N 99 CoÍelat¡on - o22 't 00 LsIFLCR Pea6on TlOSPL Peatson CorGlãl¡on N 103 N 44 - 084 LsIFLCA Peårson Coñelalion Peaßon Corelalion .053 TlOTPW 98 N N a7 - 110 L5IFRCR Pea6on Cololalion TlOIFLCA Peárson Col16lat¡on 100 N N 90 - 202' L5IFBCA Pea6on Colelalion Pea16on Colelalion 083 TI OIFFìCA 98 N N - 121 Pearson CÆrre¡alion Peaßon Corr€ht¡on - 127 FMM16 Ll M2 29 N N 112 - 063 .041 Peaßon Correlalion Pearson Colelalion FMMT L1M1 30 N N 109 - o42 . Pearson CÆrelalion Peaßon Cotelation 030 HLMl LI M6 92 N N l05 .006 HCMT Pearson Colelal¡oh Pearson CÆlelalion 087 L1M9 l'18 '108 N N Coilelalion 153 o58 FHBMl8 Psarson Ll PHL Pea6on Co(elal¡on

N 108 .052 LlPHR Paarson Corelalion '112 AGEGBOUP N -.064 045 Peaßon Colelal¡on L1M1O Pearson Coarelal¡on FLMI 97 104 N N 187 P6arson Coreialion 220' L1Ml 1 Peaßon corelalion FCMs 't't 122 N 0 N 154 Pearson Colelat¡on 158 LlSPL Pearson Corelat¡on glWM2 N 43 - 284 o ol level (2{a¡led) LlTPW Pearson Correlalion " Correlation ¡s s¡Onif¡canl at lhe 42 o 05 lev€l (zlalled)' N ' Coíelat¡on ìs s¡gnificant at lhe 063 LI IFLCR Peaßon Cor€hlion

N o22 LT IFLCA Pearson Cor€lal¡on

N

LlIFBCR Pearson Correlalion .035

N ø7 069 LI IFBCA P€arson Cþrrelation

N 94 118 L5M2 Pearson Colelalion

N 110 .000 LSMI Peatson correlalion

N 106 036 L5M6 P€arson Correlalion

N 106 ,234' L5M9 Pearson Colelalion

383 F. J. Rühti - Osteometric Variation of the Human Spine groups 8. Correlations of variables with major age gfoups separately for major time

Neolithic lBronze Age - males

AGEGROUP AGEGROUP Corelalion 459' Pearson 241 T1 M6 Pearson

N N Corelat¡on - o74 csMl Pearson Cotelation 196 T1 M9 Peaßon 23 N N Peâßon Coûelal¡on 412 c3M6 Pearson Corelalion .459' T1 PHL N 22 N 24 Pêarson Corelalion 157 c3M9 Pearson Corêlation - 172 T1 PHR N 22 N TlM1O Peaßon Corelalion -,407 CAPHL P6arson Corelation N 23 N 22 .476' f1M1t Pearson Corelal¡on øPHR P6aßon Corelal¡on 250 N N 20 TlSPL Pêarson Cotelalion 214 æM10 P€aFon Cotrelalion - 148 N l9 N 20 Pearson Corelalion 098 æM11 Pearson Corelalion - 486' TlTPW N 15 N 21 Psaßon Corelalion - 689- CsfPW Pearson Co16hlion - 315 TlIFLCB N 21 N 15 Pea6on Corehlion -.493' C3IFLCR P€arsoß Corelat¡on -,364 TlIFLCA N 22 N 21 - - 319 TlIFBCR Peãßoh Corelalion 400 C3I FLCA Pearson Corelal¡on N 21 N 21 TlIFBCA Pe¿rson Corelalion -,57't " C9IFRCR Pearson Corelat¡on - 342 N 22 N ..486' T6M2 Peaßon Corelalion 318 CAIFRCA Pearson Corêlal¡on N 24 N 22 f6M1 P6arson Corelalion 311 c7M2 Peacon Corelal¡on o47 N 22 N 22 Pearson Corelal¡on c7M1 Pearson Corelalion 125 T6M6 N N Corelat¡on 195 c7M6 Pearson Corehl¡on 560" 16M9 Pea6on N 23 N 24 P€arson Corelat¡on 119 c7M9 Pearson Corelal¡on -597" T6PHL N 24 N 24 T6PHR Pea6on C¡t€lalion .193 CTPHL Pearson Corelal¡on N 24 N 21 -.415' -.041 T6MIO Psaßon Corêlalion CTPHR Pearson Corelat¡on N 23 N -.345 . 1 Peãrson CoÍelalion c7M10 Pearaon Corelat¡on 205 T6M1 N 24 N 22 . 304 - 134 T6SPL Pêarson Corelallon c7M1 1 P€arson Cotrelatlon N N 22 206 T6TPW Poaßon Colelal¡on CTSPL Pearson Corelalion 320

N 15 Peaßon Corslal¡on 140 CTIFLCR Pearson Corelalion -.450' T6IFLCA N N 22 Peôrson Conelat¡on - 011 CTIFLCA Pêarsoh Corelalion - uo' T6IFRCA N 21 N 22 -.219 T1OM2 Pearson Corelal¡on 123 CTIFRCR PêaEon Corelal¡oh N 26 N 22 Pearson Corelalion 179 CTIFRCA P6arson Co(elalion -.467' TlOM1 N N 22 Peaßon Corelalion 320 f1M2 Pearson Co(elation ,206 T1OM6 N 24 N 24 Cotelalion .088 TlMl PeaFon Cor€lal¡on 150 T1OM9 Paarson N%

384 F. J. Rühli - Osteometric Variation of the Human Spine AGEGROUP ÁGÉGROUP

TlOPHL Pearson Corelallo¡ -.123 L5M10 Pearson Correlalion -.109

N 25 N27

fIOPHR P6arson Corelal¡on o22 L5M1 1 Peaßon Corelalìon - 183

N 25 N 28

T10M'r0 Pearson Corelalion - 435' L5SPL Peerson Coiíelat¡on - 471

N N 14

T10M1'l Pea6on Codelation LsTPW Pearsoh Cor€lalion - 310

N 26 N 12 . TlOSPL Pearson Corelalion 003 L5IFLCB Pêarson Corêlalion -.403' 't8 N N 30

fIOTPW Pearson Cþ(elal¡on 128 LsIFLCA Peaßon Core¡alion

N 17 N 28

TlOIFLCA Pearson Corelation - 319 L5IFBCB P6arson Cotrslalion 't 79

N 26 N 26

TlOIFRCA Paarson Corelalion - o90 L5¡FRCA Peãrson CorÍelal¡on - 488'

N 26 N 26

L1 M2 Pearson Coíelallon FMMl6 Pêarson Corelalion 739' 't0 N 29 N

L1 M1 Pearson Corelal¡on 098 FMMT Peaßon Correlalion 700

N 2A N I

Ll M6 Pêarson Co(elal¡on 255 HLMI Pearson Core¡alion f01

N N 24

L1 M9 Peareon Cor€lalion .206 HCMT Peaßon Corelalion - 089

N N 32

LI PHL Pearson Correlal¡on FHBMl8 Peaßon Corelal¡on 304

N N 35 LIPHR PeaEon Corelalion 369' FLMl Pearson Coûelalion 129

N N 26

LlM1O Pearson Corelatlon .033 FCMs Pearson Corelalion 't l3

N 27

11M11 Pearson Cor€lal¡on - 064 '. Corelat¡on ¡s s¡gnilicant at lhe 0 05 level (2lailed)

N 28 " Corre¡alion ¡s signilicanl at the 0 01 level (2{a¡led) LlSPL PeaÞon Cotrêlalion -,'l05

N 19 LITPW Pearson Corelal¡on .075

N I . L1 IFLCB P6arson Corelallon oal

N - L1 IFLCA Poarson Cotrelalìoh 341

N

L1 IFRCß Pearson corelalion 063

N 27 LlIFBCA PeaEon Corelal¡on - 209

N 26

Lst\42 Pearaon Corelal¡on 000

N 27

L5Mf Pearson Corelal¡on 229

N 31

L5M6 Pearson Corelal¡on .367'

N 3t

L5M9 Pearson Corelalion . 003

N s2

L5PHL Peaßon Coilelålion - o24

N 30

L5PHB P€arson Corêlallon 056

F. J, Rühli - Osteometric Variation of the Human Spine 385 Neolithic lBronze Age - females

AGEGROUP AGEGROUP Pearson c3M2 Peôrson Corelalion 038 T1 M6 N N 39 T1M9 Pearson Cor€lalion c3M 1 Peaßon Conelalion N 29 N Corelalion 062 csM6 Pearson Corelal¡on .605- f'I PHL Peaßon N 30 N Colelat¡on 021 e3M9 Peatson CorÊlalion ,211 T1 PHR Pearson 29 N N 023 csPl-ll Pea6oh corelal¡on 244 TI MlO Peãßon Coil€lal¡on

N 34 N P€arson Correlalion 112 C3PHF Pearson Corelalion 596" T1Ml1

N N Corelalion 176 6M10 Pearson CoÍelalion - 390' TlSPL PÉa6on N N .365' Pearson Colelalion c3Mt 1 P€a6on Corelalion TlTPW 17 N 36 N Peaßon Corelalion - 392' C3TPW Pea6on corehl¡on T'I IFLCR 27 N N Cor€lal¡on - 261 C3IFLCR Peaßon Corelat¡on T1 IFLCA Pea6on 27 N 34 N cor€lallon - 357 CAIFLCA Pearson Corelalion - 437" TIIFFÌCR PeaFon

N 3S N Colelalion . 038 C3IFRCB Pearson Corêlal¡on Ti IFRCA Pêaßon 27 N 36 N Peaßon cÆlelalion ,275 C3IFRCA Peatson Corelation - 460" f6M2 2g N 36 N Pearson Corelal¡on f56 c7M2 Pearson Coíêlal¡on - 171 T6M1 N N 35 Peaßon Colelalion 270 cTMl Pearson Corelat¡on - 256 T6M6 N 28 N 35 Pearson Correlalìon 085 c7M6 Pearson Correlalion 26S T6ME N 29 N 34 Pêa6on CÆrelalion 321 c7M9 Peaßon Cotr€lal¡on - 001 T6PHL N 29 N 35 Pearson Corelal¡on CTPHL Peargon Corelalion 045 f6PHR

N N Pearson CoÍelalìon - 097 CTPHR Pea6on CoÍelallon T6MfO N 28 N 33 Pearson Corelalion - 229 c7M10 PeaEon Co(elãlion - 14s T6M1 1 N 29 N 33 - 389 -.038 f6SPL Pearson Corehlion c7M1 1 Pesrson C€relalion 14 N N Pearson Corelalion .232 CTSPL P€aßon Corelalion - 208 T6TPW N 24 N . P€aßon Corelalion 451 CTIFLCR P€arson CÆrelalion - 473* TGIFLCA N 24 N P€arson Coteiation -.372 CTIFLCA Pearson Cor€lat¡on - 21s T6IFRCA N 25 N Peãrson Colelal¡on 056 CTIFRCB Pearson Conelal¡on - 300 T1OM2

N 34 N Pearson Colelal¡on l01 CTIFRCA Pearson Corelation - 396' T1OMl N 36 N 31 Peã6on Colelal¡on 106 'l-r M2 Pearson Co16lallon 057 TIOM6 N 37 N .089 TtoMg Pearson Cor€lal¡on T1M1 Peatson Corelallon 04s Nn

F. J. RühIi - Osteometric Variation of the Humøn Spine 386 AGEGROUP AGEGROUP - o11 TiOPHL Pearson Corefal¡on 011 L5M1 O Pea6on Corelal¡on

N 36 N Cotelalion . o27 T1 OPHR Pearson Corelalion 057 LsM,I1 Peaßon 40 N N ..215 T10M't0 Péaßon Corelal¡on 040 L5SPL PeaEon Corelal¡on 22 N 37 N - Corelation 079 T1 oMt 1 Pêa¡son Corelalion 351' LsTPW Pearson

N 36 N 24 198 T'IOSPL Pearson Coilelalion 148 LsIFLCR Pearson Cor€lalion 39 N 22 N - 287 TlOTPW P6arson Cor€lalion 057 LsIFLCA Pearson Cor€¡at¡on 39 N 26 N Corêlalion .. t50 T,I OIFLCA Pearson Coûelalion L5IFBCR Peaßon 39 N 35 N Corelal¡on TIOIFRCA P€arson Corelat¡on - 149 LsIFRCA Peaßon 40 N N Correlal¡on L1 M2 Pearson Cotelalion o32 FMMl6 PêaEon 12 N 41 N 12ø L,IMI Pêaßon corelal¡on 097 FMMT Poaßon Cor€lalion

N 39 N l3

LIM6 Pearson cotr€lat¡on -.009 HLMl Paaßon corelat¡on 110 38 N 3S N Pearson Corelalion 057 L1 M9 Pearson Corelal¡on 140 t-tcM7 43 N 39 N tt8 LIPHL Peaßon Corelalion ,150 FHBM.I8 Psaßon CÆrelal¡on

N 42 N - 006 LI PHB Pearson Corelal¡on ,175 FLMl Pêarson Cotelal¡on 40 N 41 N .304' LlMIO Peâ6on Corelôl¡on 't 38 FCMO Pea6on Corelation

N LIMl1 Pea6on Corelal¡on 012 " Correlalion is s¡gn¡lìcanl al lhe O 01 l€vel (2-lailed)

N 41 '. Corelat¡on js sign¡licad at th€ 0.05 l€vel (2'lailed)

LlSPL Pearson Corelation - 110 a, Canmt be coFpded bæa6e at least ore ol lhe vadables is coÑtant

N 21

L'TPW Psaßon Corelãlion 272

N l9

Ll IFLCFI Pearson Corelat¡on - 233

N 34 LlIFLCA Peatson Coíêlal¡on -,012

N 39

L1 IFRCR Pearson Corelalion - 162

N 30

L1 IFRCA Pearson Corelation 030

N 38

L5M2 Pearson Corelalion 192

N

L5M1 Peðrson Cor€lal¡on -.095

N 44

L5M6 Peareon Corelalion . 048

N 44

L5M9 Pearson Corelal¡on .005

N 46

LsPHL Pea6on Corelal¡o¡ 161

N 41

L5PHR Peaason Corelãlion - 040

F. J. Rühli - Osteometric Variation of the Human Spine 387 Medieval - males

AGEGBOUP AGEGROUP 437.. G3M2 Pearson CoÍelalion ,134 T1 M6 Pearson Corslal¡on

N 83 N 68 336- cÆM1 Peãrson Corelal¡on 156 T1M9 Pearson Corelalion 70 N N

c3M6 Pearson CoÍelalion ,394" TI PHL Pearson Corelalion

N 5S N .251' c3M9 Pearson Core¡alion ,077 TI PHB Pearson CoÍelalion 67 N N 120 C3PHL Pearson CÆrtelalion TlMIO Pearson CoÍelalion

N 60 N 61

CsPHR Peaßon Corelal¡on TlMt l Pea.son Corelalion 163

N 62 N ô2 c3Mt0 Pearson Corelalion - 068 TlSPL Psarson Corelalion 198

N N P€aFon CÆftslalion c3M1 1 Pearson Corelalion - 094 TlTFW 52 N 60 N - €fPW Pearson Corelal¡on 187 TlIFLCB P€arson CoÍelalion 002 65 N 35 N 102 CsIFLCR Pearson Corelal¡on -.045 TI IFLCA PeaFon Corelal¡on

N N - '108 CsIFLCA Pearson Core¡alion ,o24 T1 IFRCR Pearson Corelalion

N 60 N 6l

CsIFRCR Pearson CÆrelation - 0s6 TfIFRCA Pea6on Corelal¡on .o12

N 63 N 345" CsIFBCA Peaßon CorÊlat¡on 051 T6M2 Peárson Corelalion 64 N 62 N 145 c7M2 Pearson Coilelôl¡on - 087 T6M1 Pêarson Cotrelalion

N N Correlal¡on 362" c7M1 PeaGon Corelalìon ..004 T6MO Peårson 60 N 67 N Corelat¡on c7M6 Pearson Correlal¡on 57 4" T6M9 Pearson 62 N 68 N Corelalion 213 c7M9 Peaßon Co16lalion 246' f6PHL Pearson

N 66 N Pearson Correlal¡on 212 CTPHL Peaßon cortelalion 03s T6PHB 81 N 64 N i8t CTPHR Pearson Corlelal¡on 141 T6MlO P€aFon Correlal¡on 51 N N

c7Ml0 Pearson Corelalion - 035 T6M1 1 P6arson Correlalion 203' 60 N 61 N Peaßon Co(e¡ation 040 c7M1 1 Peaßon Correlatlon ,204 f6SPL 14 N 68 N Peãßon Corelal¡on 407' CTSPL Peatson Cor€lallon 0s8 f6TPW N 30 N .059 CTIFLCR P€arsoh Correlal¡on . 006 T6IFLCA Pearso¡ Cotr€lalion N 46 N ô4 -27ø CTIFLCA Pearson Corehlion - 101 T6IFRCA Psarson Corelalion 46 N 61 N 091 CTIFRCR Pearsoh CoÍelallon .212 fl0M2 Pearson Coarelat¡on

N 65 N 71 077 C7I FRCA Pearson coraelalion - o72 T1 OM1 Pêarson Co(elalion 70 N 64 N Corelal¡on 413" T1 M2 Peaßon Corêlalion 091 T1OM8 Pearson 69 N 70 N Coilelallon ,412' TlMI PêarÈon Corelalion 203 TIOMg P6arson

F. J. RühIi - Osteometric Variation of the Human Spine 388 AGEGBOUP AGEGROUP Peãrson Corelalion 204 IlOPHL PearBon Cþrelat¡on 27s', L5Mt0 N 5S N 67 * Pearson Corelalion 255' TlOPHR Peðrson Corelalion 341 L5M1 1 N 64 N 67 LsSPL P€arson Correlalion 170 T10Mt0 Peãrson Co(elalion 033 N 30 N 64 LsfPW Pêarson Corelal¡on ,256 T'toMl1 Peaßon Coß6lal¡on 136 N 2A N 67 L5IFLCR PÊa6on Cotelal¡on o21 TlOSPL Psaßoh Corelalion 314 N 67 N Pearson Corêhl¡on 076 TrofPW Pearson Corehtion 234 L5IFLCA N N ¿0 - L5IFRCR Pearson CoÍelal¡on 087 T,IOIFLCA Peaßon Co16lalion 210 N 67 N 60 't88 P€aFon Corrclal¡on TlOIFRCA Pearson Corehlion - 173 L5IFRCA N 58 N 61 - FMMl6 Peargon Corelal¡on 117 L1 M2 Pêãrson Corelalion ,060 '18 N N ø2 -.338 FMMT Pe¿rson Cþrelalion L1M'I Peaßon Co(elalion ,169 N 17 N l-llMl Peaßon Correlallon 320" L1 M6 Peañon Corelal¡on 319- N 66 N t-tcM7 Pêacon Corelal¡oh 286" LlM9 Peaßon Corelal¡on 27A' N a5 N 8t '182 FHSMI 8 P€a6on Corelalion L1 PHL Pearson Corelalion N 80 N FLMl PÊãßon Corelålion 263' L'I PHR Pearson Corelalion 114 N 68 N 78 Pêaßon Correlalìon ,2go' LIMlO Pearson cot€hlion -.00'l FCMO 69 '- al lh€ o.oi levÊl (2-lailed) LtMl1 Pearson Corelãl¡on 215 Corelalion is s¡gn¡l¡canl i5 s¡gnil¡canl.l lhe 0 05 level (zla¡led) N 74 '. Corelal¡on LlSPL Pearson CÆrelalion 22ø

N 27

LlTPW Peaßon Corelalion .308

N 27 081 L1 IFLCR Peãtson Correlalion

N 71 o52 L1 IFLCA Pearson Corelalion

N 68

L1 IFRCB Pearson Cor€lalion 115

N 88

LlIFRCA PêaGon Corelalion 121

N 66

L5M2 Pea6oh Corelál¡on

N 70

L5M1 Pearson Coteht¡on 236'

N 71 '1ô3 L5M6 Pearson CÆrêlal¡on

N 69

L5M9 P€aFon Corelalion 378"

N

L5PHL Peatson Colelalion

N 71

L5PHR Pearson Corelalion .ñT

F. J. RühIi - Osteometric Variatiott of the Human Spine 389 Medieval - females

ÀGEGROUP AGECROT,P 369++ TIM6 Peüson Corelât¡on Peffion CoFelat¡on 0ót c3tltz 64 N N ,241. TI M9 Ptrson Corelat¡on Pwson Corelalion 140 clMt 66 N N 64 196 TIPHL Peffion Colelat¡on Peüson Coreldion 395+. c3M6 63 6l N peúson Corelâtion t58 004 TIPHR c3M9 P*son coft€laLion 69 N N 6l 069 t14 TIMIO Pwson Corêlation C3PHL PersonCotreldion 56 N N 63 &relâL¡on 138 r69 TIMI I P€mon C]PTIR PemonCoreldion

N 67 -374 TISPL Pffon Corelåtion Pffion Coreldion - 2r3 c3Ml0 25 N N 50 pøson Corelation - 100 034 TITP1¡I cSMl I PeNonCorelaL¡on N 41 N Petrson CorelÂtion -l - 014 TtlFl.cR C3TPW P€ffin Corelâtion N 6l N 36 - 067 TI IFI¡A Peùson CorelaLion Pemon corelåtion -200 c3IFrcR 59 N N 6t TIIFRCR PeñonftrelaLion Pe6onCor€ldion - 244 C3IFrcA 65 N N 6t -058 TI IFRCA Pemon Corelation P€mon ColelÂtio¡ - oz9 C3IFRCR 60 6t N -m7 T6þ12 PemonCoreldion Pe6on Corela¡¡on t38 C3IFRCA 61 N N 6l - 034 T6MI Pemon CorelaLion Peüsn Co[€lâtion 104 oM2 66 N 65 339++ T6Mó Peùson Corelûtion Ptron CoFelalion 049 cTMl 65 N N 66r l4l r. PdsonCoúelütion g¡M6 PeffionColdalion 431 T6M9 65 N N 170 TóPHL Pdson Cor€lâtion gTM9 Peùson Corelâtion .31t. 62 64 N N Corelation l0l l l4 T6PHR Petrson CTPHL PemonCoreldion 63 N N 64 - 0?5 T6MIO P€60n Coreldion Petrson &relation -01I OPHR 54 N N 201 TóMI I Peilson ftrelation Peffion Corelâtion oz5 c7Ml0 62 N Corelation - 141 -ot1 T6SPL Pemon r Peilson Coreldion c7M1 16 N 235 T6TPW Peñon Coreldion Coreld¡on 125 CTSPL Peüson 4l ]J N to2 T6IFI.CA Person Cofelation Pøson Corelation -3t3+ C?IFI-CR 50 58 N N 015 T6IFRCA Peùson CorelåL¡on Pemoncoreldiotr - 096 CTIFI¡A 41 N N 51 066 T IOM2 Pemon Corelå(ion OIFRCR P6on Corelilion - 254 61 N 59 - 067 TIOMI Peõon Corelation Peffion Cor€lation -256 C/lFRcA 66 N 58 N 0t9 TI OM6 PeNon Corelalion Pemon Corelation 170 TIM2 67 69 242r Tl0I'19 PemonCotelÊl¡on TIMI Pemon Corela(ion - 040

390 F. J. Rühti - Osteometric Variatíott of the Human Spine AGECROTJP AGEGROTJP - 057 TI OPHL Petrson Corelütion .185 LJMIO Peffion Corelåtìon 51 N ._65 N 110 TIOPHR Peffion Coreldion 057 TJMI I Pefrsoû Corelåt¡on 61 N 65 N 224 Tl0Ml0 Pemon Corelation o27 L5SPL Peùsotr Corelation 24 N 58 Tl0Mu PwsonCorela(ion o47 TJTPW Pemon Coreldion -058

N 63 N 3t

138 TIOSPL Peffion C¡fi€låt¡on ú2 TJIFLCR Pffin Coælåtion

N N 64

- 006 TIOTPW Peüson Coreldion - 002 LiFr¡A Pemon Corelâlion

N 4l N TIOIFrcA Peffion Corelation 298+ IJIFRCR PeffionCoFelat¡on - 125 6t N 55 N -.201 TIOIFRCA Peæon Cor€lation t22 TJIFRCA Peffion Corelâtion 58 N N LIM2 P€ffionCorelation - t64 FWI6 P€ffion Comlat¡o¡ - 293

N 1t N

LlMI Pemon Corelât¡on - 068 FMlvl/ Pffion &Íêldion -,010

N 10 N t1

LIM6 P€mon Cotrelalion - o44 HLMI Pe6on Cotelâtion - ll0

N 66 N 54 otz Ll lvl9 Peffion corels{¡oo 144 HCIÌf¡ Pffion Cor€ldion 't5 N 69 N

LIPHL P€mon Coñelåtion o47 FHBMI 8 Peñon &ælåtion 063

N 66 N 74 - 246r LI PHR Pe6onCoftlaLion 046 FLMI Pemon &reldion 64 N 7l N 065 LIMIO Pedon Coreldion - 036 FCME Pemon Coreldion

N 58 NT d the0 0l level LIMI I Peffion corolåtion 199 'r cireldion is s¡8nilicill N 69 r Cor€lalion is s¡gnificmt ¡t Lhe0 05 level (2råil€d)

LlSPL Peilson corelåt¡on 334

N

Lltpw Petrson Corelat¡on - 57 t.*

N 23

LIIFI¡R Peùson Coreld¡on t14

N 58

LI IFI-CA Pemon coreld¡on 024

N 53 LIRCR PøsotrCoreld¡on 170

N 57

LI IFRCA Peùson coûelât¡on 056

N 56

I.JM2 Peùson Corelåtion 076

N 68

IJMI Pemon corelation 045 N 62 IJM6 Peñon Corelation 091

N 62

TJM9 Prson Coñlation 380+.

N 67

úPI Peffion Cotrelåtion o52

N 67 IJPltR Pøson Corelation - ol0 N64

397 F. J. Rühti - Osteometric Variation of the Human Spine Modern - males

AGEGROUP AGEGROUP Corelal¡on c3M2 Pêarson Corelal¡on 293 T1M6 Peaßon 3¿ N 38 N Peaßon Corelalion 438" c3Ml Pea6on Corelalion ,1 83 T1M9 38 N N Colelalion 352' c3M6 Pearson Corelal¡on 601 " T1 PHL Pearson N 40 N 36 P€arson CÆt€lalion 404' c3M9 Pearson Corslalion .580" T1 PHR N 39 N 37 Corelat¡on 20ø CsPHL Peaßon Co(e¡al¡on 468- TlMIO Peaason 38 N 35 N Corelal¡on 271 CsPHR Pea6on Corelal¡on 456" T1Ml1 Pearson 40 N N -.036 c3M10 PeaÉon Corelalion t0l TISPL Peaßon Corelallon 20 N N Correlalion 47 l" cSMl l Peaßon Corelal¡on 310 TlTPW Pearson

N N Coí€lalion - 113 C3TPW Pearson Cor€hlion 297 T1IFLCR Psarson

N N Pearson Cotelålion 090 C3IFLCR Pearson Corelal¡on - 269 TlIFLCA N 39 N Pearson CÆrêlôlion - 045 C3IFLCA Pea6on Corelalion - 065 TIIFRCR N N 35 P6a60n Corelal¡on 17ø CsIFRCR Peaßon Corelalion 124 TlIFRCA N 39 N 34 Pearson CoÍelal¡on CAIFBCA Psa6on Corelation o32 T6M2

N 35 N .073 c7M2 Pea6on Corelalion 125 f6Ml Peaßon Colélation

N 38 N Corehlion .386' cTMl Psa6on Corelalion 119 T6¡/16 Pearcon N 34 N Pearson Correlat¡on .126 c7M6 Peaßon CorElallon s87- T6M9 N s6 N Pearson Corelalion 398' c7M9 Peaßon Coilelalion 470" f6Pt-{- N 35 N 36 Pearson Correlalion 417' CTPHL Pearson corehlion 043 16PHR N 36 N 36 Peaßon Corelâlioh .200 CTPHR Peaßon Corelat¡on .192 T6MlO 34 N N Pêåßon corelalion . 059 c7M10 Pearson CoÍelat¡on .019 T6M1 I 36 N N P6arson Coielalion --41 1 c7M1 1 Peaßon Corelalion ,003 T6SPL 13 N N Corelal¡on 126 CTSPL Pearson Corelalion .,013 T6TPW Pearson N 27 N 23 Poarson Colêlat¡on 224 CTIFLCR Pea60n Corelal¡on - 200 T6IFLCA N 32 N 35 Pearson CÆrelal¡on 137 CTIFLCA Pearson Corelalion '147 16IFRCA N 32 N 38 P6arson Cotelalion .200 CTIFBCR Pearson Corelat¡on - 243 T1OM2 N N 35 Corelôlion - 006 CTIFRCA Pearson Corelal¡on 167 TlOM1 Peârson 34 N N Co(elalion 220 f1M2 Pearson Corelalion 106 T1OM6 PeaFon N 3l N 40 T1OMg Pearson CoÍelal¡on 51 5" T1 Ml Pea.son Corêlãlion 085 Ns

392 F. J. Rühl¡ - Osteometric Variation of the Human Spine AGEGROUP AGEGROUP

TlOPHL Pearson Corêlalion 105 L5M1O Pêa6on Corelal¡on - 078

N 40 N 35

TlOPHR Pearson Coilelat¡on 142 L5M1 I Pearson Coilelalion 120

N 38 N 38 572 T't0M10 Peaßon Corelalion 225 LsSPL Pearson Corelal¡on

N N Corelalion 193 Tt 0M1 1 Peãrson Corelalion 016 L5TPW Pêaßon 18 N 40 N

TlOSPL Pearson Corelat¡on 236 L5IFLCR Pearson Cor€lalion 437'

N 1l N 38 '18 068 TlOTPW Poarson Corelalion 1 LsIFLCA PeaFon Corelal¡on

N 24 N fl0tFLoA Pearson Corehlion ..006 L5IFRCR Pêa6on Cor€lallon 261

N 38 N 35 - 068 TlOIFRCA Pearson Corelalion .,086 L5IFRCA Péarson Corelalion 36 N N Corelalion L1 M2 PBaFon CoÍ€lal¡on .o63 FMM16 Pearson 2A N 36 N Corelalion - 062 L1M1 Pearson Corelalion ,329 FMMT Pêarson 2A N N .205 LI M6 Peärson Corelalioñ 091 t-trM1 Pearson Cotelallon 36 N 33 N

L1M9 Pearson Core¡alion .359' HCMT PÊaßon Corelalion 4S3"

N 37 N 40 207 Ll PHL Pearson Corelal¡on .295 FHBMl8 Peaßon Corelal¡on

N 35 N - t19 L1 PHR Pearson Corelal¡on 396' FLMl Peaßon Corelal¡on

N 36 N 36 Conelalion .275 Ll M1O Peaßon Corelal¡on 1 4Í¡ FCMs Peaßon

N 34 at the o 01 l€vel 11M11 P6aBon Cor€lalion o42 ", Correlal¡on 15 6¡gnil¡canl

N 35 ' cÆrrelat¡on ¡s slgnilicanl al lhe o 05 l€vel (2'lailed) LlSPL Psarson Corelat¡on 535

N I LlTPW Pea6on Corelal¡on .029

N 16

L1 IFLCB Pearson corslalion -.028

N 3¿

L1 IFLCA Pêarson CÆrt6lal¡on 111

N

L1 IFRCR Pøarson Corelalion t64

N 32

L1 IFRCA Poarson Corelal¡on 28s

N 34

L5M2 Pearson Cor€lalion 141

N 36

L5M1 Pearsoñ Corelalion .095

N

L5M6 Peaßon Correlalion .433'

N 32

L5M9 Pearson Corelal¡on .588"

N

LsPHL Pêaßon Corelalion 225

N 38

L5PHB P6aßon Cor€lalion 2ø6

393 F. J. Rühti - Osteometric Variation of the Human Spine Modern - females

Àoeeaoup AGEGROUP CaM2 P€arson Corelalion - 062 T1M6 Pea60n Corelat¡on 002

N26 N 27

c3M1 Peaßon Correlalion - 308 T1 ¡/9 PeaFon Coîelólion 215

N 24 N 28

c3M6 P6aßon Corelal¡on 344 T1 PHL Peaßon Corelal¡on 130

N 25 N 2A

c3M9 Pea6on Correlal¡on 369 TÍPHR Pearsoh Corelalion 224

N N 28 CsPHL Peaßon Cor€lalion TIMIO Pearson Corelal¡on 310

N 27 N

æPHR Pearson Corelal¡on 161 T1Ml I Peaßon Corrslalioh 208

N 26 N

c3M10 Peaßon Core¡al¡on 'l07 T1 SPL Pêarson Corelal¡on - 377

N 24 N 13 csMl l Pea6on Cortelallon -.'184 TlTPW Pearson Cor6lalion

N N

CsTPW Pearcon Corelalion - 493' T1 IFLCB Pearson Co16lãlion ,o47

N 17 N

C3IFLCR Pearson CÆrehtion 071 TI IFLCA Pearson Correlalion 141

N 26 N 27

CsIFLCA Pearson CÆrrelalion .008 T1 IFRCR Pea¡son Corelalion - 007

N 27 N

C3IFRCR Peaßon Corelalion - o12 TI IFRCA P€aEon Corelalion 258

N 28 N 27

CAIFBCA P€atson Correlal¡on - oo2 T6M2 Paarson Cor€lál¡on 068

N 26 N 26 -16M1 c7M2 P€atson Correlal¡on - o47 Pêaßon Corêlalion 005

N 2A N 27

c7M1 Pea6on Corelal¡on .'t79 T6M6 Pearson Corelal¡on .212

N N 27

c7M6 Pêaßon Corelalion 173 T6M9 Peaßon Corelålion - t99

N 26 N 2A

c7M9 Pearson Correlalion .256 T6PHL P€arcon Corelal¡on - 193

N 26 N

CTPHL Pearson Corelallon .167 T6PHR P€arson Corelal¡on 058

N 26 N

CTPHR Peaßon Corelal¡oo T6MlO P€arson Corelalion .348

N 27 N 26

cTMlo P€arson Cor€lãlion 324 T6MI 1 Pea16on Corelalion 183

N 26 N

c7M1 1 Pearson Correlalion - o48 T6SPL Pea6on Core¡al¡on 369

N 27 N

CTSPL Pearson Corelalion - 456 T6TPW Pearsoh Corelallon 297

N 14 N 22

CTIFLCR Pearson Coralalion .270 f6IFLCA Pearson Corr€lal¡on 181

N N 24

CTIFLCA Pearson Corêlal¡on 269 16IFRCA Pearson Corelalion 182

N 26 N

CTIFRCR PeaFon Co16lal¡on .056 T1OM2 Poarson Corelal¡on 141

N 25 N 28

CTIFBCA Peaßon Correlalion 183 TIOMI Pearson Corelal¡on 164

N N 28

T1 M2 Pearson Corelalion ..096 TIOM6 Pearson CÆr€lallon -.o21

N 28 N 26

TlMI Peãrson Cor€lalion -,239 T1OMg Pearson Corêlal¡on ..038

F. J. RühIi - Osteometric Variation of the Human Spine 394 AGEGROUP AGEGBOUP L5M1O Pearson Corelal¡on 115 TlOPHL Peãrson Cþrelalion 273 N N 2E 191 L5M1 1 Pearson Cotelal¡on TlOPHR Peaßon Corelal¡on 166 N N 27 - LSSPL Peaßon CÆÍelalion 199 T10¡¡1 0 Pearson Corelal¡on 203

N 27 -, t48 o27 LsTPW Pea6on Corelalion T1oMt 1 Peaßon Co16lalion 't5 N N . LSIFLCR Pearson Coralal¡on 150 f10sPL Paarson Correlalion osg N N 13 LSIFLCA Pearson Corelalion 189 TlOTPW PeaFon Corelalion ,214 N N 22 LsIFRCR Pearson Corelalion o41 TlOIFLCA P€aßon Corelalion 049 N 27 N 28 LsIFBCA Peaßon Corelalion 086 fr0lFRoA Pearson Corelalion 189 N 27 N FMMl6 Pearson Corelal¡on 197 LIM2 P6a6on Corelal¡on - 006 N 21 N FMMT PeaÉon Correlal¡on 122 L'IMI Pearson Corelõlion -.269 N 21 N 25 HLMl P€arson Corelal¡on - 315 LIMô Pêaßon Core¡alion N 29 N - 280 -.o75 HCMT P€aFon Correlalion L1M9 Pearson Corelalion N 30 N - FHB¡¡18 Pêarson Corrclalion 133 LlPHL Pearson Cor€lal¡oh 320

N C¡relalion is slonil¡cant al lhe o 05 level (2-ta¡led) LlPHR Pearsoh Corelalion 3't 3 N 2ø ,o23 L1 MlO Pearson Cor€lallon

N 2A

11Ml1 Pearson Corelalion 296

N 2A

LlSPL Pearson Corelalion 132

N 13

LlTPW Pearson Corelalion 003

N

LlIFLCB P€arson Corelal¡on 119

N 2A

LlIFLCA P€arson Cot€lalion -.065

N 27

LlIFRCR Pêãrson Corelãl¡on

N 27 007 L1 IFRCA Pearson Corelat¡on

N 27

L5M2 Pearson Corelalion -.144

N 26 - L5M1 P6arson Cotelalion 394

N

L5M6 P6arson Corelalion 106

N 23

L5M9 Peaßon Corelat¡on ,406'

N 27

L5PHL Pearson Còlalion 241

N 26

LsPHB Pearson Cor€lalion ,208 N^

395 F. t. RühIi - Osteometric Variatiott of the Human Spine ll I S ãCD I .t9_ .520- .o1a 42 .19 t94' .117 o qw ú2 .2a7' 23' I .076 2q' .105 o .1ú' .@9 .a20" .a76- 220' .9" 2âl' -@7 \J æHt -ø" -.239" 4 .426_ -.æô- .6 .1¡9 -.29' -.126 -.271" CD 2g- 2tT -311" eH6 ._ß1 .2$_ -ú2 .146 -210 -.@6 @5 ,os Þ o .81 .tn .329_ aM9 -.o2¡ -.ts .121 .285' -ln -.@6 -æ1 ñ 321' þ2- 276- ,723- o APHL .519' .-@o -.1S .@a .2¡6' -.ql .@7 -.oæ q .476- 236" .34- 2t3' -ffi' Ø dPHB ,9" s9" $0- -.ua --@ ,ß6_ 28' 403" æulo -ñ --on -24' -Q0 o Þ .€6_ .s8_ .370_ -.ø9 .w Æ- ,90_ .426- æiltt -3ø' -6 -.@ 219' 247. .94" .s' 272' 2g' 29. 36r .070 .sf æTPW 4' .M -197 .559- sl- .54- a .29- -20' -.130 .s_ tu @f,cF 2@' .36- .æê" 555" .727" o' .o 2æ- -w -,32- ..142 -.o2¿ .@t ætilca .5lo- 520" c) --tg -2lE' 2€ .52' .76 .7É' .lø' -0la -20' -.o24 Ø ætFce .769" .67î' --@ -.ttt -420' -497 26 -æ- I AERCA .192' -ot2 rn 3g- 230' .3@' 29" .3Ø' .38' .39- 213 CD c7m .ú9- -&- -t4l -1æ 2ê1- 267 .æ5 -l¿o -r4 -lq .7æ -tg' zl' Çil1 -.102 --@6 2fi" .æ9- -@ 26' -47 .@ .æ3 = 24' -174 -gt- 239' a- c7il6 .@ -.û5 --M o \ .92- 2f 216' --@ -137 æ cNg .107 .024 242- 25 .ú2 -.oa5 -14 -411- .æ .6 -tt2 Þ CTPHL 311- 2g' .u5 .lll --16 -N 2¿6- 2U- -tT .æ- -87 .t1t -110- F CTHR æ- .t$' CD pt5' 28" -95" --@5 .tta 5q- .4¿- -.025 Ø cilt0 .tð -.o4 --æ 2¿T' 4T .170 262" -.1 l9 .æt .39- -61- -tn o' CilII -læ ,Ét -.M .6t (D .o71 -.o12 .N ..174 -207 -ffi -.N -,@5 qSPL -.tE -lt7 .516- -49" -.12 --lÉ .&- .3tt -471" cnftcR .tlt .@2 -231' ,ø { .6r' .81' .s¿- .N -.t71 --ol5 .&" .46- .06 .&- p- cnftu .119 .@5 --l3l ,ga' .g- -99- .M --16 -.u9 -62- .52r 265' -519" Þ CNFRCR 218' .w -2&' -a7a' .ffi- -4m" -gT' 0q -.u7 -.on -.01 I -81' 4tT- CilFRCA -175 .@9 --ll5 o .319' 233' .la2 -ts' 2ç' 236' -423" 4- -3&- Ttw -.0fi .99- g4- 24' .136 .36- .É .123 -04 ftut .3%' -Æ .171 .-ø3 -.ot2 -.075 1M 27 Æ' .# .t6 ,02 2g' TIH6 Æ- 2t0' fr- .ø4 .u2 .og .117 CD .302' 216' 2g' -.136 .lg .326- flM9 25' .16 2U- .N .6 --@ .on Þ .18 36" fl- .l5l .t05 .319' TIPHL .39- 3m- -3øt- --o1a --@4 .0F .4d- .1Aê .& 2g -.ot1 -175 2U' .131 34' È æ- .u2- --N .lt3 -@- -49' -tt9 371" 3ú' .@6 --o76 ø2 TIMlO 2g' 1q -t2t 2q' .010 .l@ 259' .fl- æ TIHII 23' .ut -ot5 lri -149 Þ .tg .tol -14 .*- -.ñ .ta5 .ø .8 Tlsil c)

(¡) \o o\ S 4il2 qut æM6 æM9 æPHL qPHã qUlO æN1t æfPw æIFLæ æIFLCA æIFRCF AIFFCA

I fIIFLCF -197 .@ -.t5t -t71 -.04 -.@7 .3q- .321- -ù2 .392.. .3¿6" 342- o ntftca .ua -.16 -.140 -.@o --tg -.1ø .a97 3V -6t .3S- .fl7. .45" .M -.od -lto ..@6 ..070 .g- t¿0' -1ú 3A" -fl' .4n.. -423' a fIIFFCE -r4 nlnca .t2a --l07 ..æ9 .@ -.tt5 -.019 .tft- .olt t4" &" -F9' ì f6M .97' .@- 2' 249- .96- .2tt' .197 .t7' -12 -tÆ -.o70 .12t Til' .tt5- .12 235' 4' .121 .ltl .96" -.61 -ot9 -.14 .Mt ñ f6H6 3m' .t% t4' .159 2tt' .t4" 24' .82 .tt¿ .oa6 -@ -.@ -.N5 I ftw 34' -tt7 246' g4' 4- 2¿r ..o11 ø' ,430" .o29 .tæ -ßo -135 96" .t27 26' .tn- .127 -o19 .lM -.N o 16PHL Jn- .t4a 2A' 211' -1F- .4t' .tn 21t 2ta' -3@é .1n 2T .no' -76 .l2t .@a .o Í6U10 -.t07 -26' --ut 230' -.1ü .-æø 4. .37v -,M -146 -137 't-lgt .t97 T6U1l -1fl -.ú6 -.61 .t9t .67 .on 5la* .@5 .1at .lâ6 .tB -1tt f6sil .tæ .û7 .o1t -27 .att .@a .,1t6 -2É -t9l -.@l .la6 .t29 -ts f6PW -on -.at 2t .t5c Æ' I .tt9 .u7 fl --M --@9 --M -.@5 T6lf,CA 230' .87 .ofi .132 -.1 lo --N .om N' 2¿9 .359' 4' .291', .3@- tr Îdæ4 .tß .@1 -.N .@4 -.113 -.1@ 2aø 2fl. -04 .39' .32¿' .3ú- ftow -s7 -w .0n .1t9 2Þ* 212' 2a7 .@7 222' .o27 .1fi TIOHl .!f- 2T .oß .M .tæ .løa ,r1ø -176 .tu .É2 .117 .Øt -.012 ftoM6 .371- .tt7 2tt' .lß .tt6 .2¿9' ,t02 24' .127 .0^ .l@ .M -67 ftoB9 -lt6' tttè 27r. .1æ # â5- -.@ .36' 39" .o5l -18 .167 .230' r'OHL 4' .æ1 -121 ,t8' n20a 2t' .ts .æ1 -t% .M .N ftonR 3nr ,36- .æ 2lî .!f9- 259' 2n' 210 .052 .187 .od -@7 ftoüto -.M -.tlt --æt .ot I -.14 ,llt .te .w' 24 2@ 210' -lta 235' Tloill1 2fie .oã -.019 -lt9 -.119 -oF 36- -96- .34' 274' &7' -319" flOSPL .@1 .120 -.116 .il2- .t@ ,û' .tß 2æ 21 .1â3 .t& .@2 .@t -.Éo 233 21t 2t .176 âl .093 .on .ß .N ftolf,4 ,@ -.67 -2¿r -.121 -2t6' -210' zl -.033 .ffi' .18" -49' ,519" notna .o8 .-l3t ..t8 -.@9 -2U -,ot9 286. -476- -.oÆ .tg- .45'. .419' LIM2 JA- .127 .tg .90- 2&' 26' .w -.ot2 .04t .87 .og -o4 Llul 5N' .t@ .t2l 2il' ,læ 2fl', .Æ -1il .l@ .02ø -.@ --o12 LIN6 ,1rc ä2- tl3' 210' 2' .127 .t69- .013 .w -.d1 .@o LIM9 .ß2' 246' .ttJ" 269à 2a3' 24' .w' .ût -05t -.ß -67 LlPHL -92" 2n" -l¿¿ 26' .&- ,36- Æ' -û4 .162 .M .110 -.@ LIPHB .39' 2a1' .131 .4F .39- .tt0 .ot2 .67 -o73 .122 .ot9 LlHlO .16 -.@7 --@a .tg -.60 .o75 ,37a- .44- .360- .tu ,tn

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È co TlOIFRCA LTM2 \ TlOSPL TlOIFLCA T10M10 floM11 T1N6 f10Ms TlOPHL TIOPHR S TlOM2 TlOMl à _.w .361" .285- 295" -,178 1S -.073 - 079 .223' -.o16 ..172 271_ o52 .on .@ -.o14 .1r0 37A" I .126 2a5" -21A' .19 .æ7 .119 .2Æ' 029 I .02 .@a ..231 .147 ,gt' TlIFLC -ql -.q6 272" .259_ .os ..€7 ,365- .345" .09 .o10 -æ2 .2æ" .089 .079 TlIFRCR ,@ 1a7 .g- 151 .t10 .ß9 - olr .97" .to1 6 .1n .t2á TIIFFCA s7- 395" .155 g - oga ..1G7 .s_ .a33" .au" @ø .!2!' -,o59 T6M2 .t71 .@3 ì -2É' .212' .051 .193' -42' .n4' 479- .224 .125 TsMI .a21' .12¿ .173 .&r -.66 -.124 .39" a71" .l¡7 .694" g9' .o25 Til6 .g- .@3 d7 -7ú" .@_ .o15 336" I -271' .s2_ ,a29' .69 .æ5 l6M9 .45_ .113 .æ5 .3S" a$" .a$_ -o51 .313" -læ .ag_ .q ,G7 T6PHL .sr .1a2 .033 s$'r a .426" 528" .$5_ ls7 æ1 .215" g_ .107 .211' f6PHB .a55" .s2a_ .s" .13¡ lal .217' 124 .135 .19¡ .18 , -a .1$ .1€7 -on 217' T6M1O .121 .558" ,æ2- 251- .076 -120 .142 .-125 21' -@2 476' .102 -.031 T6M1I 220r .1m -.078 .@3 .os 4@" .1@ ..1æ .@ .07{ .@ .a$" .ta5 qa T65PL .371_ .216 .¡a3- 235 -29" .2¿a" 030 .117 -.d ,0a1 .@1 TôTPW .197 .19 .on .6 ,142 .259' ..64 .017 .39" .076 252' TsIFLCA 6l @ .ts .2@ .0æ .tq .ll6 516- -.to tg 075 -21 ¡' .q3 aâ T6IFFCA .92s" .271' &' .@5n .s7_ .ó? 4fl" \ .315_ .0tl ,075 .2S_ .262" .@" now .v0- .za2' .1r?' .152 .145 .3S_ ,070 @0 .6n- 06 107 -412" froM1 -297" ,3S" -316_ 3$_ æ3 - 071 -.1 10 flow 8' -1tî ,53" .2lG' .2ll' .s5_ .051 @ 3q- .Ø- .N' .3&" .1S .26ó -.@ Ttoug -276' -w .295" .1a1' .2al' .0lr -.@ T1ffiL .618- .w- ,211" .112 g- 279" t74 -6r 24- -on 26 .&a" -97- I .89_ -14 .ti7' -t@' ,209- .q .302- .234' .6i !17' .265' floilto 267 -al4- -9f, 8" -.o21 .@ 2*' 2g -1U -tl -2Sl ñoult .@5 239 -l¿2 .64 -161 .o5_ .42" 1$ .l# Æ .lq 3n' f1osil -.øt .16 -126 .127 276- .o52 -u7 I 741- ÍtoDW --117 .M â7 .u1 .o4 -.o7t --æt .a@- .G8 -.0m -ttø .@1 .61- I ftolftd .æ9 .ñ 97' 316' .61 .014 .lg .æt ,ß -.M 1 .lû .on .195 .og --orc rtolfrca -361" .ol6 .ñ .182' -39" -.@5 .79' -gf -4r -317 2¿2' .ot2 --@o LIM2 2&- -tl4' Jlø" -.û5 .@7 .92' .fr' 39' .-o* 2l' -l& -le -&* LIMI .1n- 210' .lc9' .w .124 26' 28" 8a' .169 --llt .012 LIN6 .M #' 223 )6' .É" 28- .167 -.16 Æ- .16 -6f -lN .æ5 -.o8 Llw ,162" tz- .@7 .ltt -lg' --16 --19 52' LIPHL 2V -3æ' -.67 215 .139 .11¿- -tn* --18 271- %' 24' .ffi -3À' ,tt0' 4t' -6- &' _i36 .127 .ñ LIHIø 4' .ø'

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2æ_ 234', ¡¡5_ .171' ,216' .198 .225 ,oa7 .125 I c3M2 .3S_ .29t" .242' l@ .212' 2S- .162 017 162 1æ' c3Ml 314" 95" .2a5_ .42" .s5" ..@ .@o .274 471n -.1 e2 -.1a0 6A $ w6 .82 .ß5" .515" 1g .222' ,046 - -.120 ,g -.@5 .@ .22a' .s9.. .03 -.@f 2g 2r7 c3M9 1S 1S .2tt' .183 .6 N .16 ß .!79_ a7a" .@3 ..m2 .!15. -2a7 CIPHL .1t3 .a79- .2Aî' ì -.1n 014 .os .105 .320_ ,ß1" ..@l .015 227 .t$ o C3PHA .120 &" 22' .245" at1" .110 .61 127.. 3$n ,270' Ã\ dt0 q0 169 . t10 .0$ -ú7 .129" æf .65 .s_ .31a" .!tt_ .253' ì 312_ -1ß .2lt' .g_ .s_ È c3Mlt .215' .165 ,204 371_ 226 .363_ ,312' .415- 67 .21€ .124 C3TPW ,29' .2t' 224 o .-m .2S- .1æ ,295" -o7 -o12 3n" -12a .æ9 -2æ'r , ÉIFLCF .ß ..os -.071 ,G .252 .225' 230' 110 .2æ' .@Ê ,ffi ,29S" .146 -.0s c3lFLC^ 220' .@7 .lt5 . .90" .121 .929- .@o gð_ ,d - otl t53 .29' €IFRCR .tg -,207' -.@ -.0t2 .265_ .20' .0¡4 tq -.@7 2{1' .1{ - o5a .@ c3tFac^ .073 .09 .@5 .oa ..M .M2 217' .159 .121 -272" .201' 24" .214 c7M2 370" ,2øT' .329* .3ta_ .@4 .l$' .@ .1a7 226' ü2 .114 .90' -.013 c7M1 -s_ .217' .259_ .€- .t& .l ó1 .o52 .3tt- .269 .252 .G5 .@ b c7M6 ,@l .as_ .s1" .¡12_ -2U" .G7 -o71 \- 28' 65 0?5 ..07t .107 -.t6 .035 c7M9 .115 .s- ,a73_ 2ae' .@ G2 -.t11 .171 @t .120 .2A! .lza -2!' CTPHL .115 -251' -28' .19' ..121 -1a7 .t73 .@ .232' .26_ .@ .l5l ,171' .1lt GTPHR .2@' .!23- .2Sn .s8- 251. $0" -.125 .s_ .31t- .12C .æ 3q_ qM10 -121 -.1@ .0tl -g -.t73 6 .20!' .193' .123 ,189 .129 .19 ,079 .370'. ,2a7" cTMl 1 ..115 @6 t93 ,1S _æ7 ..q .$9i 49_ .@1 ,62 .124 CTSPL 29' .@- .46- .t¡l -.@6 .142 ..0a1 .t29 ..1Þ -.99 .212' .t@ qFLCR -@3 ,ot3 ..142 .,107 .r10 .427" 28_ $1_ @7 .36a" .172 @1 .044 .398" cTlFLC^ .G .o$ ,of2 .67 ..51. -019 216' ,039 .o32 -61 ..161 ..117 -.q .2€1- .19 CTIFRCR -.0s -.on .12.- .366" .1æ .t l6 .14 .92" -a20" .211' cnFRc^ .1al -62 .d5 .ø1 ,@ - 69 .69 .t55 .215" .152 .t6l .2æ .1æ tta TIM2 3$- .370" -2î' .t80 .163 .217' .150 .z7a .g' .1N .1S' TlMI .511" .211' ,29' .211' .268" 210' 017 .zao- .2â1 .1rl @o .11t .tg ltMs .M .175" .a76_ 2a3" 2W' .@a 1t7 .1æ .17s -245ù 01t .079 -g ,142 T1M9 -o7a .2!t- .2€5_ -29' 03Ê .gs .olo .2æ'. -.12! .w .s' .tg - laa TlPHL 29" .z1T .295" 270" -.145 .ql 071 .246_ .o35 ..w 267 .t1! ,ø6 TlPHR .170 .152 -224' .tæ. .329- .12Ar. .0t .521" a0_ .o71 1tr .27â' as2' TlMlO .g 1S .168 ,@ .237' .i@ -L1" .33!- ..09 ..159 .232' .25t_ 213' TIM'I -.oto .ot .149 .2@' .oa6 .o17 .1 s7 .072 29' .121 @1 .113 2gl .4S' TISPL .l4l 3n" -221

rÞÊ H LTIFFCA L1M11 LlSPL LlIFLCS LlIFLCA S L1M9 LIPHL LlPHR LIMIO à LlM6

- 159 013 .,og .126 .5rl- .l3l .127 I I LISPL 3æ' .lô6 ..o$ ta6 222 o LIPW .fs 217 -.6 --w .tt1 .lat 2 z4 t .æa" .570- LIIFLæ -ot7 .M -.otl -.tg --161 3E' .tB" -& .la7 I o .7s1- Lilla .tat .N --107 -.01t -.o70 .59" .374- .14 271' ,s6' t LllncR -û6 .ßo -.ola --@5 -.@7 .tÐ' 2g' .tæ .llo -nl' .52' I ì 57 I o LIIFRCA -1ø1' .tß -,og -.@7 --02 -6tr .171" .t1t 2U 597' -t@" -.039 .o76 .o74 .lØ' Lsw .3æ- 227 .t4- .tgÈ .0F 217 -.M I .@ø -.o26 .tg L5Ut ä0' .34- .!si .3@" 210' .3@- .læ .tt9' -û1 2ß' 24' -.ot0 .N .ú .Èl ..0m -.@o -.olt -og o w6 -a2a' Ð -.1 .M 213' L5ß 34r .&' -97 .t1r 3B- .83 27f .tta t7 -175 -@' -Õ .@t .JA' .ûê -.110 .og -.o41 .1 t¿ EPHL Jtt' .ß' 2S .ß2' -tt2 238 -,8 -ol3 --M .@ LffiR 26- 2ê9á 3A- iu- .1U ,o52 ,go- .s6' 27 -lØ 24' -.o16 Éi Lil1O .æ3 -.u3 .oF .16 -.o23 ø 2a1' 3*" 2ú- LsM'I .ll9 4l' 29' -lat' .16 .&r &" 8' 45- 2tt -.162 -.014 .02 .M ßSPL -t9 .Jæ. .301. .w N .oz --Æ -4t- .230 .lt3 226 Lmw .N .ü7 --æt --on -.6 .om .æt .@ .lø .tt2 €" .174- EFLCN .B ,o8 .0ru .t01 .on .tto" .3Ø- .96" -M .&1' -.@1 351 - 2ß' ,374- Ãftca .B ..on .og -.@i -ú7 -321" -39' Æ7 27¿- --û9 .81' .322' gncn .on -012 .o41 -t4 -,og .a2t- 260- .tl6' .352' ßFed -ta7 --ot7 -t9t' .69 -0fi .JF .t9r n .1n t26'- .tß .N .lø ilHl6 --tu .1æ .N .E .o2a 26 231 --út .tt1 2g -271 .330' .84 .la6 3A' ffi' .7ß .tv ñu7 -.11 I -ø --û5 -.133 -69' .t3a -tD HLUI -aæ" .34* .17- tr Æ" 4' .3U- tu 2l 4r .w Hffi t6' 24- 36' .121 .12' .&' -13ø -w -.ût .ot5 .ot7 -.oÆ ffi" FHBNI' -ar9" -tu" 4' 2* ß1' .3¿2' a7' -i04' .ll2 -tn lHt .475- 276' tfi- 2&- 29" 239" .121 .t07 flt' .lat .ot4 2V 274' .@ .@6 --oÞ .@a FCgC 2n' 3F aD- .191' 43È

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È (JlÉ L5IFLCA EMlO L5M1 I ÉSPL 5PW LsM9 L5PHL ÉPHR S sM2 L5M1 L5M6

øil.l.th ..0¡5 .t@ -.1t .436r ..03 @ 1$ .111 -276' .q6 .232 .1æ .320' -.146 -.o72 -@3 I LISPL .032 .g' -1@ ..070 .æ5 .110 za2 -.140 -.of2 .37!' .!g- ,329" .313- s2" .147 -29' .qt .@5 ..075 -æ7 -.0s7 -.032 420" L'IFLCR É .133 .3t!' .39" o ..@7 .o3a -297- 2*' .102 .@t .o71 .i62 45" LlIFLCA .16' .295' .g' ,463_ .176 .@o .@2 034 .93 .m .214' .@6 ,99- .(9- -a11- : LlIFRCF .2t5_ .29_ ..@ .174 .141 .@7 .170 .læ .ls @7 4 .Qa -.t20 -o3l LlIFFCA .s7_ -.172 -.124 18! .1at .126 .32f ..1S È\ ,3S' .¡16_ .,1þ -.i43 LsU2 .o1" t56 .65 .æ" ì 21tr .2€1_ ..qo -29" .19' &l .4r -.q0 .16 .2{ .01! .€9_ -2ã7' .d6 .sæ- .t9 -2¡0' .o32 -125 ' a &6 .21t' .@ .26" .@3 .037 M 3æ' .141 .210 .1S -.111 .o w9 .6S- ..09 .ü7 .tT .@7 -gt .117 úilL .42' .,G .107 .191 .146 22r 42- ,ü9- .32a" 191' E*A -2fl' -tt7' ..o72 .175 .1ø -Ø7 -@2 .ñ -224' --@ Ë -.o74 270' -17 .339- ßNl0 41" .369' .ø7 .339- .1a7 ,qa .60 EUlt -.É2 .o75 -47 -tÉ .w .126 -.161 .65 -32t' -136 .@ .201 .@0 fsPl --ø, ,17 .lt .019 b -.@6 .otl -u5 -ß2" .7@- -.@9 Æ -.@l .212 \ mw -.o75 4v .3%- --N1 .ore .@ 418_ 6ftCR .139 .tt5_ .ov .41à ..745 --@ .013 .ß -.ß .123 .o4 -agr -170- ßf,Ça .mt .3@' .&f -4 .tlt --t@ -op -ñ -flg" gncÈ -ol7 .tÑ- -.og -o7a -471- -747 .& --a1 --@ -111- -.@ 4' --62 217 -131 .2ft Étnca 2û .l$ -.lsl 2ø 212 ø -1t7 .116' -Nl -.ttl -. 234 -26 ilut6 .26 371' .64 f8 -07t -.61 .t8 --M --u2 ,u1 .137 .l& 26' FNNT 213' 2tt' .t1r -.094 -t52' -167 251' 3g¿ -tl3' -0lo .@7 .N HLNI -49' .tæ .110 2rê' -.@ .3@- .l¿4' 2Ø' .t23' .3@- -la4' -lco' ,tF -lt5' rcN7 .tæ .r3!- -ats" -.011 -t70' -&' -3ú- .tt2- ilEUlt .tq' -.o17 .w -t3l -tt7 Æ- 2@' .t@ 21ø' -lg -ú9- äl' 2T .16 -tt9 -tæ' Fø1 tÉ 2g- -sil' --æ7 2t' 4l' -7a9' 2l' FCW st'

IF F o\ >! FLMl FCMs atw2 ! FMMlô HLIú1 HCMT FHBMI8

tu.l.tld tur.htld .€7_ .40" 291" $2" 357- eM2 ..t41 .95' ,215 .ru- I .1æ .2a1 ru- ,24' .a@" ,s- \) c3M1 -.1æ .1t7 .051 .@2 .s!- 322r .@3 .s7_ c3Mg -.219' .217 0f5 o ..@5 al_ 2S' -.æl .2S" .@ €M9 - 170 .97' -.070 -523' .3!- .117 ru- ,29 EPHL -.117 .1$ .g -1ry .@ .1r0 ,a36" ,2S- .ls .27a" Õ BPHR - 162 .113 .?42 ..1 1t .N .s0' .1S - 076 .æ? 07t 5\ c3M10 .29_ -215 .a69- .210" .s_ .s3- .æ1" -aa" æM1r -.0æ 9T $t' 334_ .325_ ..m9 gfPw .q .41 .æ1 .314' ,a10" .ß9- o .É q -,@ -.61 .øE dFLCA .240' -.@ -.d5 .073 .o 055 ..0tt .u2 .61 .26' øFLCA úl -010 -on ,24' .ot 211 . .1@ -.@ @5 ,øa C3IFFCR .220' 01â .os .92.. .211' ,67 -.w -.M -,@ @FACA .2@ .161 .@l E .2&" .156 ,325_ .176 m2 ..1G .2U .151 -2a9', .289- 22a' .2S_ .@9 079 .zl3' 46' .24" Ë cTMt -.os .160 2S' .go_ -137' -2a1' .313_ 224 (- @M6 -,8 ,24 .@7 g 279' .145 -27i' .s_ .r45 .s- m9 -.161 .111 -.014 .t33 .1q .115 .1â5' .201' .139 CTPHL ..074 .@ .1V .m ,q9 .llo' ,255" .053 .tg .0$ CTPHR -ø0 -.61 .2U' .075 .1@ .16 .o7a @Ml0 .1U .aæ- ,2a2', ,103 .29' .t05 -2â7' .1 87 .1& .2a7 c7M11 .tg .2ø .371- -219 .æE .1S 4$" øSPL -.@ 2X9 .m .o11 .w' ..@ .1æ .1t a ..É -.071 .g CNFLCR 230' .ßg 220 .2S .,Qt .ta€ 1g .053 ,@6 æ3 cTlFLC^ Q5 26 .,07! -013 -@ .,@8 1@ ,28r qIFBCR .151 .1æ .371- .fæ .on .0t .19 øFnc^ .ln .@' -.013 2€' .155 .224 zaa-' .2E9" g" .320" ,s_ T1M2 - 19' ,173 .2U .2S_ -261- -1ø7 182 .276" ,21t -w' T1Ml .,6 .189 .s9_ .112 .@ .1@ .s9- a15" .13 T1 M6 ..65 3n" .136 ..ls .186 245_ ,39" -2n" -27' TIM9 -.H .æ3 -.o21 .@ô .26!- 26{_ M 287' TlPHL ..lg .132 -67 .215" -æ7 -.07t .126 251_ 2ú" .148 TIPHA ..o74 .127 .2S' .207 .185' .!s .138 B T1MlO 13S .223 -203 .117 .o1 3 .153 .1 6? 217 .262 .118 ,078 T1M1 1 .125 .@ 376_ .É- .24 .375" .s' TlSPL - 235 .19 -lg

È Ê\ :: S HWI HCMT FHBMl¡ FLMl FCMs EtWM2 à LsIFACA FMM16

..073 .!51 .272 ,112 ..@7 ..w -.03 I .24' .gg' .059 .! 15 .226 O TlIFLA .t& 231 132 .16 129 .@¡ ..@ ..@7 -w ..d5 1$ TlIFACR .ts .gl' .121 os a .u7 .ql .61 2a7' tl¡mc^ ,20' .149 .1ß ls ,m .ß9_ .aGz_ .a70' ,a8ô- .16t ì Til2 -.@1 .172 .16 .284- 211' ,s" .s7- .sl_ ,103 T6MI .æa . o7t .82 .11G .47' .€1_ g7_ .3æ'. .159 TôM6 -.2@' .261 -.01t s_ .491" 210' .9" 267 ì -.18 .110 .oa5 -2n" @- È T6M9 .g- .go- .276- .gg- .ts T6PHL ..t35 .2aa ,tq .ær o .166 .214 .s5_ .sf .sg- .95_ .9" T6PHR -.1c0 .9" ..m .o 120 .la0 .la2 .21f' .1f1 T6MlO .q€ .0aa .127 -01a -22¿' 075 .112 .2S T8Mli -,@a 43 .45A" .14 .@2 -.122 .017 -.1æ .@ -.@1 TCSPL -.01 I -.ffi -.66 .39- .a&" .$9" .g féPw -.t23 .@ .g .H_ .297' ò .107 .245' .ß2 .4 T6IFLCA .olo .o1 6 .6 .@ -.@5 .059 0a! -.u7 .@e 3?' b T6IFRCA ..@5 .N .165 \- .97_ .&- .sr' .395- .l1t' T1il2 --o75 .t tl 270 -a22- .@ .G" .175 2S_ .2ø7' TlWT --g - 121 .2Ú .2S' .511_ s6" .ffi" .466_ 24' 110M6 -.t47 .2s1 .@ ,gt- 2S_ .44" .a13_ .272" -aT" 204 TIOM9 -ls .2ff 110 .193' 2a6" .317_ 1æ .æt- .ll4 TlOPHL -.@ -.o54 .119 .232' .$3_ .291_ .tl 2¡9_ .170 TIOPHR .,61 ..0æ .18 .201' .214' 207' .t62 275r fIil10 .138 &3' .32A' .2la- .116 .16 .w .115 .210 ItNll -on .316' .95' .221'. ,310' .524' .320' .12 .a6" .166 T1oSPf -.1S 207 401 ,364_ ,19 ,235 .23i. .248 TlOTPW .63 .274 .014 ,312" .m .t6l .111 .æ9 .317. IrdRc^ .171 .164 .H .2t6 .G5 6 o72 .@l æt TIdFRCA .193 æ3 ..151 .167 .2E!_ 322_ .370_ .429- -323' L1M2 ..tt7 .217 .m ,&" o71 .214' .1& 2s2- .19 LlMI .ol 8 .07a -.1@ -2311 31a" .@_ 122^ 25t" .9" 2at LIM6 . i70 .ß5 o72 .21!- .{0" ¡39" .29" .aa' .s' L'M9 -.19 .o71 .o27 .a@ .14t .s1" .g_ 201' .3S- 075 LI PHL .@t .1 li 070 .1@ .9" .107" .19 .275' LIPHR -.u7 .@ -.ls ,232' 0g -216. .i63' .o' LlMI O -255" .æ5' .ø' J¡"" * "" "'' "^'

ÊÈ @ +1

FHBM18 FLMI FilE S L5IFRCA trM15 HLMl H47

-l7a -,24 .051 ,t25 .@- I .@2 .€ .128 .175 .272 .221 .1S 101 o LITPW .æa .qt -0$ ..N .142 ,o71 2*' .2S" .æ. @ .2æ" .m5 o LlIFLCF ..@7 .@o .157 .1S' .G .os LlIFLCA .s2_ .1ß 23' .323' .,@ .110 .0s7 .119 1m .@ LlIFRCA .195' -.87 .€t .171 .ta2 .1$ 16 LItFAC^ .Æ- G9 .on .219' .0s ..6 .317" .320- 2n' .2¡lÚ ÉM2 -.19 .1S -25¡ .492- .s" .@" slù \ uMl -.1n .26 .017 .s5.. .261" .a17' ,52a' .43!" iær .fl2" LS6 .3S" .211 ..N .s_ a ,&1" 27ø" .G- -.o12 sl -.172 .39' .@5 ,!73- m- -.@ -o -.211 .2S" .517- .215' 29" L5PHL -N - lta .29" .3æ" ,ts .,1€ ..193 .2tr_ ,?Ul u- ,21f LsPHR ..071 0t .2A' 127 .1al .m sr0 .Ð- .132 .015 .@5 .152 .14 -221' .29- .2S9- 1trl1 .28" -337' .!$ .276" -.021 .@ .ß' ..t46 -211 .2ú' ESPL -.087 -.1S .og .310 326' .210 .{2' ,39- -2ST lâ LfPW .075 il5" 279 .,1{0 2n' -.9 .073 -@5 0s LsIFLCR 324- .63 -67 ,os -.016 -ts .lÚ -.1æ -on EIFLCA .705- .ú2 .1S .1æ .029 -.æ1 -.o11 -0t0 HFRCR .181 -u7 .E -.on .231' -l{ ..0æ ..1t -,151 LtrFA @ 24 .31¡' .97' .15¡ .2S .037 FUUI6 2& 224 .ø1 -.rc .,q3 ..051 .12ø FUHT 27 ;g- .sa" .w- .813_ .s1_ .42- HLUI 272' & .1lo .$e- .416_ Hil7 .lu -216 -.ln .624" FHEUI' 259' .lß 2& -7ßñ .€- .s2_ .67v FW 2ll' .1n .tn .n7 .4t" 91- g- -ffi' FU ,la -6 .u7 .g- -gs-

-. @d.ttú t.dFtbñ¡.t h.o.ot bd(2ud)- 1 M.ilù I. JilltuI.t tu ùÉ Ld (+blu).

rÞ \o 1 0. Signif,rcant microevolutionary regresslons Variable (m ales) Method r Signif icance

Log o.77 0.00 agegroup Log 0.48 0.00 TH10 spinous Process length 0.45 0.00 THG leÍt caudal intervertebral foramen width Log 0.00 bi-iliac width Qua 0.44 0.43 0.00 C3 spinal canal transverse diameter Qua o.42 0.00 TH10 vertebral body sagittal diameter Log 0.00 C3 left cranial intervertebral foramen width Qua o.42 0.00 C3 right caudal intervertebral foramen width Qua 0.40 0.39 0.00 C3 left caudal intervertebral foramen width Qua Exp 0.39 0.00 L5 transverse Process width 0.00 TH6 sagittal diameter vertebral body Exp 0.39 0.38 0.00 THG right caudal iniervertebral foramen width Pow 0.37 0.00 L1 spinal canal transverse diameter Exp Log 0.36 0.01 L5 sPinous Processus length 0.00 femoral head width Exp 0.36 0.36 0.00 humerus length Qua Log 0.35 0.00 THlO transverse Process width 0.35 0.00 C3 spinal canal sagittal diameter Pow 0.34 0.00 L1 left craniall intervertebral foramen width Exp Log 0.34 0.00 C3 dorsal vertebral body height dorsal 0.00 TH1 spinal canal sagittal diameter Pow 0.33 Pow 0.33 0.00 C3 transverse Process width 0.00 L1 right cranial intervertebral foramen width Log 0.32 0.31 0.00 C3 ventral vertebral bodY height Pow 0.00 L5 left cranìal intervertebral foramen width Log 0.31 0.00 TH10 leÍt caudal intervertebral foramen width Log 0.31 0.00 TH10 right caudal intervertebral foramen width Log 0.30 Pow 0.30 0.00 C7 right Pedicle height 0.00 C7 left cranial intervertebral foramen width Exp 0.30 0.00 TH1 right caudal intervertebral foramen width Qua 0.30 0.00 TH10 dorsal vertebral bodY height Lin 0.29 0.00 L1 vertebral body transverse diameter Qua 0.29 0.00 C7 transverse diameter spinal canal Log 0.29 0.00 TH1 spinal canal transverse diameter Pow 0.29 0.00 C3 right cranial intervertebral foramen width Exp o.28 o.28 0.00 L1 vertebral body sagittal diameter Log 0.00 humerus circumference Log o.28 0.00 L1 left caudal intervertebral foramen width Exp 0.28 0.00 C7 vertebral body sagittal diameter Log 0.27 Pow o.27 0.00 TH6 right Pedicle height 0.00 L1 spinal canal sagittal diameter Log 0.27 0.00 THl left caudal intervertebral foramen width Log 0.27 0.00 C7 dorsal vertebral bodY height Log o.26 0.00 C7 right caudal intervertebral foramen width Pow 0.25 Pow o.25 0.00 TH10 right Pedicle height Pow 0.25 0.01 C7 left Pedicle height 0.01 C7 left caudal intervertebral foramen width Log 0.25 0.01 L5 spinal canal sagittal diameter Log 0.24 0.01 TH6 transverse diameter vertebral body Exp 0.24 0.01 femur length Lin 0.23 0.01 femur circumference Lin 0.20 -0.33 0.00 C3 vertebral body transverse diameter Qua 420 F J. Rähti - Osteontetric Variatiott of the Huntan Spine Variable (females) Method r Siqnif icance

bi-iliac width Cub 0.55 0.00 C7 transverse diameter spinal canal Cub 0.51 0.00 C3 right cranial intervertebral foramen width Cub 0.49 0.00 TH1 spinal canal transverse diameter Cub 0.47 0.00 L5 spinous processus length Log 0.45 0.00 C3 spinal canal transverse diameter Exp 0.43 0.00 0.43 0.00 f emur circumference Cub femoral head width Cub 0.42 0.00 L1 spinous process length Log 0.39 0.00 Log 0.38 0.00 TH10 transverse Process width TH10 right caudal intervertebral foramen width Qua 0.35 0.00 humerus length Exp 0.35 0.00 L1 right cranial intervertebral foramen width Cub 0.34 0.00 TH10 spinal canal sagittal diameter Cub 0.33 0.00 L5 left caudal intervertebral foramen width Log 0.33 0.00 TH1 0 vertebral body sagittal diameter Log 0.32 0.00 TH6 sagittal diameter vertebral body Exp 0.31 0.00 TH10 spinal canal transverse diameter Exp 0.30 0.00 L5 right cranial intervertebral foramen width Log 0.30 0.00 L1 spinal canal sagittal diameter Log 0.30 0.00 C7 left caudal intervertebral foramen width Lin 0.30 0.00 C7 right caudal intervertebral foramen width Lin 0.29 0.00 Log 0.01 TH1 transverse Process width 0.29 C3 spinal canal sagittal diameter Log 0.29 0.00 C7 right cranial intervertebral foramen width Log o.28 0.00 L5 left cranial intervertebral foramen width Log 0.27 0.00 L1 right caudal intervertebral foramina width Log o.26 0.00 TH't0 vertebral body transverse diameier Exp 0.26 0.00 THl right cranial intervertebral foramen width Log 0.26 0.00 L5 right caudal intervertebral foramen width Log 0.26 0.00 TH6 dorsal vertebral bodY height Log 0.26 0.00 TH1 spinal canal sagittal diameter Log o.26 0.01 L5 spinal canal sagittal diameter Pow 0.25 0.01 Ll left caudal intervertebral foramen width Log o.24 0.01 L'l spinal canal transverse diameter Log o.24 0.01 agegroup Log -0.15 0.04 L5 ventral vertebral body height Pow -0.26 0.00 L5 vertebral body transverse diameter Qua -0.28 0.00 TH6 right caudal intervertebral foramen width Qua -0.37 0.00 femur length Cub -0.39 0.00 C3 right caudal intervertebral foramen width Qua -0.44 0.00 C3 leÍt caudal intervertebral foramen width Qua -0.45 0.00 TH10 leÍt caudal intervertebral foramen width Cub -o.47 0.00 C3 left cranial intervertebral foramen width Oua -0.56 0.00

427 F. J. Rähli - Osteo¡netric Variation of the Human Spine 11. ANOVA of variables with major time groups (Non-Bonferroni tables: bold= significant; italic : decrease)

s¡s

f10M6 11 22 000

I r 0tu19 192 151

TIOPHL 253 oð4

IIOPHR 262 077 Males T10M10 344 035 fl0Ml I 414 018 TlOSPL 693 oo2 Sig TIoTPW 417 019 AGEGBruP s42 oos TlolFLcÀ 626 003

c3M2 693 001 flolFRc 735 00!

c3M1 413 018 LlM2 11 897

c3M6 101 566 LlMl 71 192 833 OoO LfM6 007

C3PHL 459 012 L1M9 487 009

C3PHR 313 048 L'IPHL 628 002 c3Mf0 s32 006 LIPHB 340 036 c3Mll 754 001 L1Ml0 016

C3SPL 14 ajz LlMll 707 001

C3TPW 317 o{8 LlSPL r20 308

CsIFLCR or4 L|ÌPW 60 554

C3IFLCA 562 Oo5 LI|FLCR 390 023 C3IFRCR 182 167 LIIFLCA 196 145

C3IFRCA 681 oo2 LlIFRCR 124 016

c7M2 172 184 LI|FFCA 227 L5l!12 cTMr 91 4oi 164 197 c7M6 oo1 LsMl 67 513 c7M9 876 L5M6 340 037

CTPHL 198 142 L5l!19 54 586

CTPHR 3 3t o4o LsPHL ,27 766 c7lvt0 95 .39r LSPHR 08 927

c7M1 I 617 000 LSMTO 405 ,020

CTSPL 259 oø2 LsMl 1 198 t4c

CTTPW 193 162 L5SPL 765 00t

CTIFLCR 96 304 LsTPW 153 226

CTIFLCA 392 Oz2 LSIFLCR 7t6 001

cTtFRCFt t63 2o1 LSIFLCA 4A 018 CTIFRCA 592 004 L5IFRCR 143 .243 trM2 35 708 _l-5 FMM16 805 4s3 TlMI 25 780 3 675 032 T1 M6 310 049 HLMl 15 t82 000 T1 M9 60 550 HCMT I 693 000 TlPHL 195 147 FHBMl8 14 412 ,000 TI PHB 46 631 FLMI lo 215 000 T1MlO 488 009 FCMS ô 450 OO2 TlMII 736 001 ðililr¡. / <ñr n! a fl SPL 203 138 206 t33 si9 TlIFLCR t68 .191 F 9 074 000 TlIFLCA 443 ot4 C1M6/M9

1 9fi 406 IIIFRCR 33g C3M1 o/Ml 003 IlIFRCA 037 c7M6/M9 6 1s7 2 012 138 T6M2 424 Otl C7Mlo/Ml 1 I 784 173 T6MI 01 991 flM6/Mg 436 648 f6M6 t0 39 oo0 TlMlo/Ml1 I 980 000 T6tú9 158 zto T6M6/M9 I 461 632 T6PHL 147 234 T6M1 o/Ml 5 703 004 T6PHR 385 Oz4 Tf0M6/Mg 155 457 f6Mlo 330 041 T1 0M1 0/M I 1 L1M6/M9 €93 502 T6MI 1 l.3s 262 086 918 T6SPL 622 11M10/Mr I LSM6/M9 2677 o73 T6TPW 28 760 L5M10/M1 t 4 360 015 T6IFLCA I66 000 FMMT/M16 i 880 164 T6IFRCA 815 oot 5 904 004 floM2 11 47 ooo HM7/M1

F. J. RühIi - Osteontetric Variation of the Human Spine 422 sis

TIOM6 623 ,003

T1 OMg 1 ,71 185 IlOPHL 106 350 TlOPHR 1 41 249

Females T1 0M t0 578 o04

T10M11 428 0t6

11 0sPL 239 101

F. s¡s TtoTpw 709 o01

AGEGROUP 290 .058 TlOIFLCA 914 000

c3M2 226 109 TTOIFRCA 876 000 526 c3M1 65 L1M2 c3M6 66 518 LlMt .79 454 466 011 c3M9 L1 Mô 85 .431 199 CsPHL 164 Ll M9 s8 .417 482 C3PHB L1 PHL 22 802 269 073 c3M10 L1 PHR 70 496

c3M1l 11 60 .000 LlM1O 489 009 C3SPL 500 011 L1Mll 424 016 C3TPW 356 034 LlSPL 492 .o11 c3tFLCÊ 810 00'l LlTPW 295 081 001 C3IFLCA 772 Ll ¡FLCR 15S 210 127 E3IFBCB 210 L'I IFLCA 215 121 o23 C IFRCA 389 L1 IFBCB 6.S2 .003 1ô8 f91 C7M2 L,I IFRCA 298 .055 820 cTMl ,20 L5M2 o7 933 038 c7M6 s35 L5M1 285 .061 83 440 c7M9 L5M6 97 381 423 CTPHL a7 L5M9 343 035 135 .263 CTPHR L5PHL 45 636 c7Ml0 1.54 218 LsPHB 35 703 000 c7M1 1 18 78 L5M1O 266 .o7 4 182 CTSPL 175 LsMI 1 168 190 315 CTÍPW 120 LSSPL 462 014

CTIFLCFì t57 212 L5TPW 195 150

CTIFLCA 339 .o37 L5IFLCR 004 o57 CTIFRCR 294 L5IFLCA 713 001 CTIFRCA 368 028 LsIFRCB 605 003 q2 T,I M2 93 r srERc . .ì?, - .054 ,947 T.IMl 24 .785 FMM16 FMMT 3 033 o57 T1 M6 80 453 HLMl 5 666 .004 T1 M9 10 906 HCMT 2 931 0s7 T1 PHL 782 FHBMIs 13,548 000 T1 PHR 13 874 FLMl s 083 .000 Tl MíO 333 039 FCME 15 072 000 T1M11 14 67 F'WM' lffi fISPL 227 TlTPW 393 o23 Sis. f1 IFLCR 145 240 F 2.7 45 068 T1 IFLCA s5 .s77 csM6/M9 483 6t8 T1 IFRCR 254 o83 c3M10/Ml'l 4 581 .o12 T1 IFRCA 106 350 c7M6/M9 4728 01'l T6M2 476 010 c7M10/M11 .l09 T6MI 341 TIM6/M9 .694 4 544 0t3 T6M6 727 001 T1Mt0,/M1t I 327 oo2 T6M9 71 493 T6M6/M9 948 T6PHL 98 380 T6Mt0/M1 1 't.356 262 T6PHR 1.ø7 l5s T10M6/Ms ,999 371 T6M1O 155 ,217 T10MtO/Ml 1 636 531 T6M1 I ,a7 420 L1M6/M9 724 447 T6SPL a7 .42ø 11Ml0/M1',| 1 062 349 T6TPW 96 388 L5M6/M9 I 044 356 T6¡FLCA 124 289 LsMr0/M1 1 3 ô01 036 T6IFRCA 033 FMMT/M16 13.o53 000 T1OM2 .61 545 HM7/M1 w T1ôMr 'i ¡d ffi FM¡/M 33rO = 423 F. J. Rühti - Osteometic Variation of the Human Spíne Multiple Comparisons - males

Bonferon¡

Mctn Dependenl (l)ljre (J)ljæ D¡llêrerce Deperdenl (l) ïìæ (J) ljm O¡fererce sis Variable group gror+ (t-J) Var¡able grow 9roup (I.J) sig 100 C7M9 2 -17 AGEGROUP -30 .o7 100 3 - 50' 00 CTPHL -40 ..20 34 -34 22 c3M2 2 100 05 100 3 -,87' 01 CTPHR -40 I ,00 3 - 46' .04 caMl ..38 47 -36 91 - 81' D2 c7M10

2 -43 18 15 100 c.3M6 -24 100 -1 tl o8 ,48 c7M1 1 3 -2 14' 00 3 100 'I -t 03' 04 c3M9 2 85' 00 70 't.00 3 'I 16 CTSPL -1 79 59 3 -69 24 -2.48 08 C3PHL 2 55 09 -14 90 18 -02 100 CTTPW 3 -1274 47 2 - 56' 03 2,16 1.O0 CsPHR 21 100 -21 t00 -24 83 CTIFLCR -35 50 2 - 49' 04 t,r 100 csM10 -60 s7 06 -1 32' 01 CTIFLCA 3 - 96' 02 3 o9 -..17 '1 00 csMl l 2 -1 24' o0 .03 100 3 -1 47' 00 CTIFRCR -3l 46 -23 to0 2 30 €SPL -63 100 - 89' 02 -91 100 CTIFRCA 3 -1 14' 00 2 -,24 100 -25 100 csfPw -2 19 24 2 -23 100 -3 36' 04 T1M2 -27 100 2 '1'16 .86 2 -05 L00 CAIFLCR -48 26 -25 100 . 88' 01 TIMI 'I 00 -.41 23 'I 3 08 00 CsIFLCA - 86' o4 -.63 31 -1,23' 00 TIM6 ¡ 07' 04 2 -.37 58 -44 56 G3IFRCR - 31 ,60 36 '1 00 -50 't8 T1 M9 2 3 1.00 2 l9 t00 -61 2 3 ,86 CsIFRCA -,85' 03 56 19 3 -1 31' 00 T1 PHL 56 25 3 -15 31 3 .00 1.00 c7M2 -32 67 -05 100 .53 20 fÍ PHR 2 18 1.00 -21 100 23 100 cTMl - 14 1.00 -60 10 100 T1MlO 3 - 94' 0t 3 36 54 -34 ,46 c7M8 2 -t 01' 01 -85 19 -1 47' 00 TlMIl -1.83' 00 -.46 40

c7M9 - 15 100 a'9ß

424 F. J. Rühli - Osteometric Variation of the Human Spine Mean Mean Deperìdent (l)'tìre (J)'rìm Dilererce (l)'tìre (J) Tìm Dìfererce (l-J) sis Dependenl Vadablê grcup groUp grouP (t-J) sis Variabþ 9roup 100 -43 1.00 T1OM2 TlSPL 2 .81 06 3 -2 41 TIOMl 3 -60 2 ¡98 26 3 100 -308 26 fITPW i -2 50' 00 -3 75 15 TlOM6 -3 46' 00 3 -Ê7 100 -.96 33 .36 47 tt tFLoR 2 -1 19 21 110M9 -t 46 2 - 14 100 -27 100 1.00 3 TIiFLCA 19 -62 20 97 .04 TlOPHL -79 09 o3 1.O0 .05 100 17 T'I IFRCR 1.00 -31 67 TlOPHR 2 't0 -28 55 -47 -48 46 2 T1 IFRCA -69 -9t' 03 T10¡,llo - 90' o4 -43 3 -21 100 -1 04' 0l T6M2 2 2 ¡ 04' 03 3 -86 10 f10M11 -l 't0' o3 Í8 100 -,05 100 -01 1.00 2 T6M1 -2 11 03 100 TlOSPL -5 93' 00 2 3 .o4 100 05 -2 00 T6M6 20' -2 42 38 00 TîOTPW 2 -4 90' 02 3 100 -2 4A -90 2 T6M9 -88 12 -69 65 TlOIFLCA -1 63' 00 2 3 21 100 3 -75 14 .40 51 T6PHL 2 -,63 42 OIFBCA 3 .53 T1 -1 71' 00 .,13 100 ¡ 08' 02 18 3 TsPHR -09 1.00 02 L'IM2 2 3 06 1.00 2 ,60 2 100 2 -.17 100 f6M10 100 3 07 LlMI 't1 36 100 49 71 -06 '1 00 T6M1 1 -1 09 27 -53 .54 Ll M6 .2 35' 01 3 ..47 .1 26 155 100 T6SPL -1 76' 03 100 LI M9 -2 26' 0l 100 3 2 69 2 - 5'l 100 -t 08 100 T6TPW 09 100 3 -1 33 100 LlPHL .83' 04 3 100 00 'I 00 T6IFLCA ô3 -2 14' 00 LlPHR 2 -32 '1 00 00 2 .1.69' 70' 04 -.09 1.00 T6IFRCA 2 2 -76 08 3 .1 40' 0! L'rMl0 -1 09' 0l -1,3'l' ,00

-1 64' 00 T1OM2

425 F. J. Rühti - Osleometric Variation of the Human Spine M€an Mean Deperìdenl (l) lìG (J) Tire D¡flererce Deperdent (l)'ljre (J)lìm D¡fererc€ group gfoup Variable group gfoup Sg Vãriable (t-J) sis LI -1 10' 01 LsIFLCA 3 -'t1 100 3 00 L5IFRCR -07 100 -36 .43

't 2 42 L1 SPL -30 00 -2 69 ,ß L5IFRCA -29 100

-2 3S 51 - 19 100 'I LÍTPW -2 17 1,00 2 1t 00

.4 57 88 Êl,lM16 -l 08 86 -2.40 100 -1 15 a7 2 -07 100 L1 IFLCR -47 - a2' 02 FMMT -2 25 08 ..35 -2,45' 03

LIIFLCA -52 56 -88 15

-36 99

'|00 Mean L1 IFRCR -05 Dependenl (l)'tìre (J) liæ Diflererce -67 05 Var¡able group grc|lp 0-J) sis 2 - 62' 02 HLM1 -21.13' 00

L1 IFRCA 2 .44 -1234' 0Í

3 -91 't1 3 -8 76' 03

2 -36 92 t-tcM7 -3 12' 00

L5M2 2 .51 79 -4 46' 00 s 18 100 3 135 39

2 3 .70 28 FHBM1 8 -2 95' 00

L5M1 -43 100 -3 35' 00 3 -69 75 3 2 39 1.00

-26 100 FLMl 2 -26.55' 00

L5M6 18 3 -15,60 06 -f 96' 03 3 -.l0.95 12

-72 .42 FCMs -4 70' 00

L5M9 -98 .91 s -3 23 10 -65 1.00 3 -1 47

'l oo BIWM2 -9 23 31

LsPHL 00 1.00 -16 88' 0f

25 100

23 100 ' The rean difererco ¡9 s¡gn¡f¡canl al the .o5 level

L5PHR -01 100

3 14 't 00

3 .15 100

LsM1O 2 -51 09 -1 47' 02

2 3 -.96 11

L5M1 l -1 12 3 -86 50

2 i00

LsSPL 194

-3 21 12 -5 t4' 00 LsIPW -44 1,00 -8 14

-7,70 34

LsIFLCR -04 100

3 .01 - 66' 00 'I LsIFLCA 2 ..35 00

426 F. J. RühIi - Osteomelric Variation of the Human Spine Multiple ComParisons - females

Bonferon¡

Mean (l)1re (J).Iìre lJlf ererce (J) D¡tlererce Dependenl Dependenl (l)rìre Tru group grol+ (¡-J) s¡g grouP (t-J) Variable Variable -48 c7M9 2 2 -21 91 ..34 11 CTPHL 03 100 09 1,00 -24 ø2 -26 3 c3M2 2 - 31 33 -59 11 CTPHR 3 -26 76 -04 100 'I 2 c3Mt -11 00 .45 35 -37 ,78 c7M!0 42 26 1.00 08 100 c3M6 2 90 .79 2 -1 47' 00 3 26 100 c7M1 1 -2 64' 00 2 o4 100 1 17' 0l 02 csMs 1 26' 1.73 20 129 o6 CTSPL 121 90 3 .03 1 00 100 39 3 2 -52 C3PHL 29 2 -10 62 ,41 00 100 CTfPW -4 A4 1.00 2 29 47 -5 98 1.00 .06 100 3 CAPHR 2 12 1.00 - 16 100 CTIFLCR 3 - 4'l 25 68 3 23 2 54 c3M10 1.00 -58 13 .09 CTIFLCA - 83' 05 .59 't9 2 23 1.00 -1 04' 00 3 c3M11 2 2 .,17 100 -1.58' 00 CTIFBCR 3 -53 05 54 ,24 36 22 1Z 100 CASPL -59 09 -3 03' 02 CTIFBCA . a2' 04 I 2 92' 24 100 100 2 C3TPW -1 02 -35 62 -3 17', .04 f1 M2 -38 215 13 03 100 00 3 CAIFLCR I 2 f 2 1,00 94 o1 T1 Ml 1 00 'I .22 s 02 00 - 10 100 -1 00 CAIFLCA 10' -30 | 00 .1 25', 01 fl M6 3 -.47 .65 06 100 2 17 100 -45 24 æIFRCB o8 1.00 -57 TI M9 27 100 ,12 'I 00 .19 'I 00 - 03 3 c{il FRCA 84' -17 100 't3 2 3 -75 f1 PHL -07 100 -06 100 3 100 .43 10 c7M2 -03 100 -31 a7 T1 PHR 'l00 3 09 3 ..12 I 00 2 100 08 100 c7M1 47 TlMto 3 19 1 ,00 -,ø2' 03 '! 2 00 2 43 38 14 c7M6 2 -1 61 00 - 88' 05 flM1l -2 30' 00 3 .30 100 100 c7M9

427 F. J. RühIi - Osteometric Variation of the Human Spine .tìm Oêpendent (l) lìm (J) Diferêrce Dependenl (l)-t¡re (J) Îru Dillererce Sg (t-J) sis Vañabl€ group Var¡ablê group 30 88 TlSPL 2 100 100 -1 8l 63 110M1 2 1l 244 1l 3 74 63 2A TlTPW -2 80 13 -1.05 425 02 T10lvl6 00 L45 69 -09 100 .88 T1 IFLCFI 2 -36 35 TIOM9 58 3 -1.06 2 27 ,48 2 43 f00 - 31 100 3 T1 IFLCA -03 100 .38 100 TIOPHL -45 60 o7 100 100 41 56 T1 IFRCR -02 -45 21 fIOPHR 21 100 3 .30 100 3 2 .43 11 -?4 100 3 2 51 30 T1 IFRCA 2 2 -.33 77 3 -57 45 T'0M'10 3 .1 17' o0 2 98 03 T6M2 -24 1.00 -64 12 .1 03' 0l T10M11 -1 06' 02 3 79' 03 4l 66 T6M1 -35 100 -01 100 flosPL 2 3 .2.75 13 3 2 33 ,70 3 2 240 22 T6M6 2 -1 55' 00 -2 30 22 s -1 96' 00 TlOTPW -5 5t o0 40 't 00 .40 .06 T6¡¡9 -1 ,00 .05 100 TlOIFLCA 20' -1.20' -00 -35 1 00 3 .o8 1.O0 T6PHL 2 21 1.00 -.s4' o1 -07 100 11 0lFRCA -1 41' 00 2 ,24 63 -.17 100 T6PHR .48 18 LtM2 82 100 46 24 '1.00 '1 00 2 38 T6M1O 41 80 -53 25 L1 MI 2 02 100 3 2 29 01 '! .13 100 39 00 T6M1 f 2 -08 100 -53 .64 L1M6 -72 6S 40 8g 3 2 64 73 T6SPL t,95 a7 -60 76 100 L1M9 2 -04 74 220 100 .,90 '1 00 17 T6TPW 2 16 100 -216 .52 L1 PHL 1.00 126 100 3 05 100 69 3 16IFLCA -52 31 81 -7ø 38 LI PHB 07 100 2 25 100 100 03 24 T6IFBCA 2 -t 03' .,40 -44 100 LlMlO 52 -1 09', 01 ..60 41 100 TlOM2 02 3-nffi

428 F. J. RühIi - Osteometic Variation of the Humnn Spine Mean Mean Dependenl (l)ïre (J)Îm D¡ffererc6 Dependent (l)-t¡æ (J) Îre Difererce Variable group (t-J) Sg Variablê groç 0roup (t-J) Sg LsIFLCA 71 11Ml1 2 1B .l 19' o1 L5IFRCB 98 -.88' 00 56 44 02 LlSPL 60 100 .,37 92 .3,31' 04 L5IFBCA 2 -1 20' 02 3 3 S1 ,01 3 2 .83 LlTPW -2 03 100 2 -35 100 -6 24 o6 FMMl6 -29 100 2 421 ,29 100 3 2 -06 1 00 L1 IFLCR 11 .28 FMMT 2 .19 1,00 4l 44 -1 69 21

-.1 100 LI IFLCA 4 .15

3 58 .24 Mean (J) D¡ffererce Ll IFRCR 56 11 Dependenl (l)ljm lìm (t-J) Variab16 gror.P group sis -38 68 HLMl 2 -12,44' 01 2 93' 00 -12 E3' 02 -28 96 L1 IFBCA 39 t.00 -82 05 t-tcM7 89 7A 54 -1 20 65 'I L5M2 o6 00 3 20s 05 -.12 100 -1 00 FHBMl8 2 96' 17 100 -2 56' 00 -.1I 100 L5M1 .59 82 3 -1 45 o7 FLMl -20,57' 00 127 12 2 -17 43' 02 08 100 L5M6 -3 09 'I O0 91 58 -4 00 FCMs 44' -83 63 -6 50' 00 L5M9 ,54 100 3 206 2 20' 03 -11 BIWM2 84 3 2 -1 66 12 -22 21 00 2 30 100 LsPHL 3 , 11fl4 'I 3 42 00 ' The Ean difererce ls sign¡ficanl at th€ 05 level 2 100

LsPHR 00 1.00 100 -33 'I 00

LsMIO -.54 75

3 -1 31 07

2 .77 42 -,78 46 L5M1 l ¡ 14 27

36 100

L5SPL -,l.30 68 -3 84' 01 254 14

L5TPW -t 85 100 -9.35 ,18

745 34 't.00 L5IFLCR 11 3 01

2 .68', 01

L5IFLCA 2 - 06' 05 3 -156{

429 F. J. Rüht¡ - Osteometric Variation of the Human Spine i2. Alterations of standard deviation of variables with time group I versus 3 (F-values) Variable (SD) - males F (Time group 1 versus 3) SD'Ti SD - Time qroup 3

agegroup 1.40 0.7 0.8 * c3M2 2.32 0.9 1.4 c3M1 1.08 1.0 1.0 c3M6 j: 1.58 1.9 1.5 c3M9 1.17 2.0 1.8 CsPHI 1.00 1.1 1.1 C3PHr 1.05 1.0 1.0 c3M10 1.10 1.5 1.5 c3M11 1.63 1.3 1.7 CSSPL 1.90 4.1 3.0 C3TPW 1.23 4.2 3.8 ** C3lFIcr 3.17 1.5 0.9 G3lFlca 1.33 1.5 1.3 C3lFrcr 1.25 1.1 0.9 CSlFrca 1.15 1.4 1.3 * c7M2 2.01 0.9 1.3 ** c7M1 2.87 0.9 1.5 * c7M6 2.33 1.1 1.7 c7M9 1.42 2.7 2.2 CTPHI 1.10 1.0 0.9 't.0 CTPHr 't.02 1.0 c7M10 1.10 1.5 1.5 c7M11 1.06 1.8 1.7 CTSPL 1.22 3.9 4.3 CTTPW 3.51 9.8 18.4 CTlFlcr 1.25 1.0 0.9 CTlFlca 1.34 1.3 1.5 CTlFrcr 1.73 1.0 0.7 CTlFrca 1.53 1.5 1.2 T1 M2 1.34 1.2 1.4 't.1 T1M1 1.37 1.3 * T1M6 2.12 1.4 2.0 T.IM9 1.35 2.4 2.8 T1 PHI 1.10 1.2 1.3 TITPHr 1.18 1.2 1.3 T1M1O 1.02 1.3 1.2 T1M11 1.15 1.9 1.8 T1 SPL 1.44 4.3 3.6 TlTPW 1.52 6.3 5.1 Tl lFlcr 1.40 1.2 1.0 Tl lFlca 1.00 1.5 1.5 T1 lFrcr 1.14 1.0 1.0 T1 lFrca 1.10 1.4 1.5 T6M2 1.24 1.5 1.4 T6M1 1.77 1.3 1.7 * T6M6 2.46 1.6 2.4 T6M9 1.65 1.8 2.3 T6PHI 1.45 1.5 1.1 * T6PHr 1.94 1.5 1.0 T6M1O 1.18 1.2 1.3 T6M11 1.42 1.6 1.5 T6SPL 1.27 6.1 5.4 T6TPW 1.55 5.6 4.5 T6lFlca 1.72 1.3 1.8 TGlFrca 1.49 1.6 1.3 F. J. Ri¡htí - Osleomett'ic Varialion of the Human Spine 430 1.5 T1OM2 1.26 1.3 * 1.5 T1OM1 2.03 1.1 * 3.3 T1OM6 2.44 2.1 * 3.4 T1OM9 2.O2 2.4 1.4 TlOPHI 1.35 1.7 1.2 TIOPHr 1.41 1.5 * 1.6 T10M10 2.03 1.1 1.8 T10M11 1.63 1.4 3.1 3.7 T1 OSPL 1.42 TlOTPW 1.00 4.9 4.9 Tl0lFlca 1.18 2.O 1.8 1.00 2.0 2.0 Tl0lFrca '1.6 L1M2 1.44 1.3 2.2 L1M1 1.32 1.9 2.9 L1M6 1.10 3.0 ** 3.4 L1M9 2,60 2.1 1.3 1.3 L1 PHI 1.11 * 1.1 'l.6 L1 PHT 2.28 1.7 L1M1O 1.50 1.5 2.0 11M11 1.62 1.6 ** LlSPL 4.66 2.8 6.1 LlTPW 9.12'* 3.8 11.5 Ll lFlcr 1.18 1.2 1.3 ** Ll lFlca 2.93 1.4 2.3 * Ll lFrcr 2.31 0.9 1.3 * Ll lFrca 2.39 1.3 2.0 1.9 L5M2 1.17 2.1 2.3 L5M1 1 .18 2.1 3.1 L5M6 1.61 2.4 * L5M9 2.23 3.4 5.1 1.6 L5PHI 1.20 1.8 '1.9 1.9 LSPHr 1.10 2.3 LsM1O 1.60 1.8 3.0 LsM11 2.21 ' 2.O 3.9 L5SPL 1.48 3.2 ** 7.6 LsTPW 5.09 17.2 1.0 LSlFlcr 1.O4 1.0 * 2.3 L5lFlca 2.17 1.5 1.0 LSlFrcr 1.16 0.9 1.7 L5lFrca 1.O2 1.7 2.6 2.6 FMM16 1.01 2.3 FMMT 1.55 1.8 HLMl 1.41 14.1 14.5 5,6 HCMT 1.26 4.4 3.2 FHM18 1.18 2.8 25.3 FLMl 1.05 20.9 6,9 FCMs 1.23 5.8 16.6 BIWM2 1.20 10.8

bold increase (signif icants onlY) italic: decrease (si gnificants onlY) significant before (P<0.05) / after Bonferroni's correction

431 F. J. Rühli - Osleometric Variatiott of the Human Spine qroup Variable (SD) - females F (Time qrouP 1 versus 3) SD - time 1 SD - time orouÞ 3

agegroup 1.74 "" 0.63 0.83 * c3M2 2.19 0.91 1.35 c3M1 1.15 1.13 1.21 c3M6 1.12 1.33 1.41 c3M9 1.02 2.34 2.37 0.93 C3PHI 0.99 0.94 C3PHr 1.58 0.78 0.98 * c3M10 2.07 1.03 1.48 c3M11 1.81 1.18 1.58 C3SPL 2.92. 2.37 4.05 * C3TPW 2.58 2.44 3.91 C3lFlcr 1.45 1.31 1.08 C3lFlca 1.65 1.61 1.2s * CSlFrcr -2.08 1.51 1.05 C3lFrca 1.67 1.71 1.32 0.97 1.41 c7l,!12 2.10 " * c7M1 1.87 1.02 1.40 c7M6 1.58 1.19 1.50 c7M9 1.30 2.12 1.86 0.97 CTPHI 1.20 0.88 1.08 CTPHr 1.40 0.92 1.34 C7M1O 1.79 1.00 c7M11 1.43 1.49 1.78 2.86 CTSPL 2.10 4.14 21.79 CTTPW 2.22 14.64 CTlFlcr 1.07 0.82 0.85 CTlFlca 1.23 1.18 1.31 CTlFrcr 1.03 0.83 0.84 CTlFrca 1.11 1.11 1.17 T1M2 1.31 1.21 1.38 1.40 T1M1 1.38 1.19 T1M6 1.46 1.32 1.60 2.06 T1M9 1.06 2.00 1.27 T1 PHI 1.41 1.07 TITPHr 1 .16 1.16 1.08 1.18 T1M1O 1.65 0.92 1.68 T1M11 1.57 1.34 TlSPL 1.04 3.08 3.02 TlTPW 1.81 5.24 3.90 Tl lFlcr 1.97 " 0.72 1.01 1.34 Tl lFlca 1.21 1.22 0.92 Tl lFrcr 1.13 0.98 Tl lFrca 1.34 1.33 1.54 1.40 T6M2 1.09 1.34 1.17 T6M1 1.11 1.23 2.31 T6M6 1.61 1.88 1.96 T6M9 1.36 1.68 1.25 T6PHI 1.70 0.96 1.20 TGPHr 1.85 0.88 1.02 T6M1O 1.14 0.96 1.75 T6M11 1.08 1.68 s.64 T6SPL 1.13 5.29 T6TPW 1.39 4.99 4.23 T6lFlca 1.14 1.84 1.97 T6lFrca 1.14 1.80 1.69 432 F. J. Rähli - Osteomett'ic Variation of the Hr'unan Spine 1.75 1.45 T1OM2 1.46 1.54 1.75 T1OM1 1.30 1.93 2.47 T1OM6 1.64 2.54 2.35 T1OMg 1.17 1.38 1.70 TlOPHI 1.52 1.43 1.68 T1OPHr 1.39 1.39 1.30 T10M10 1.14 1.54 1.71 T10M11 1.24 4.30 4.13 T1 OSPL 1.09 5.01 4.73 TlOTPW 1.13 * 1.12 Tl0lFlca -2.21 1.67 1.15 Tl0lFrca 1.44 1.38 2.04 2.04 L1M2 1.00 1.88 1.84 L1M1 1.05 2.07 2.64 L1M6 1.63 2.92 2.80 L1M9 1.08 1.23 1.36 L1 PHI 1.22 1.38 1.53 L1 PHr 1.23 1.22 1.55 L1M1O 1.63 1.55 1.77 11M11 1.31 3.30 3.99 L1 SPL 1.46 6.53 LlTPW 1.15 7.01 1 .15 1.37 Ll lFlcr 1.41 1.55 1.45 Ll lFlca 1 .15 1.12 1.30 Ll lFrcr 1.35 1.31 1.35 Ll lFrca 1.05 2.24 1.79 L5M2 1.55 2.54 2.08 L5M1 1.48 2.66 2.68 L5M6 1.O2 3.11 3.22 L5M9 1.O7 1.87 2.23 L5PHI 1.42 1.77 2.O9 L5PHT 1.40 2.38 1.90 LsM1O 1.57 2.75 2.93 15M11 1.14 3.21 4.06 L5SPL 1.60 * 16.72 10.56 LsTPW -2.51 0.94 1.14 L5lFlcr 1.84 1.82 1.64 L5lFlca 1.23 *' o.71 1.28 L5lFrcr 3,26 1.88 1.58 L5lFrca 1.41 2.74 3.49 FMM16 1.62 3.00 2.76 FMMT 1.18 ** 17.29 21.16 HLMl 1,96 ** 4.14 5.01 HCMT 2.10 ** 2.34 3.17 FHMl8 3.30 *' 23.37 25.27 FLMl 1.80 ** 5.46 6.09 FCMS 1.93 17.17 15.69 BIWM2 1.58

bold: increase (signif icants onlY) significant before (P<0.05) / after Bonferroni's correction

433 F. J. Rühti - Osteomelric Variation of the Human Spine present and selected 13. partial correlation coefficients of variables with time before long bone measurements (variables with significant correlation with the selected long bone measurements only; whole sample) Maximum femur length:

- r Variable - males Variable females

* 0.16 c3M2 0.22 c3M2 ** 0.30 ** c3M1 0.29 c3M11 ** 0.38 ** c3M11 0,27 C3IFLCR 0.33 ** C3TPW 0.15 C3IFLCA C7M2 0.05 T1 M2 0.14 c7M1 -o.14 T1M1 0.03 ** c7M11 0.37 T1 M9 0.15 0.06 TlTPW 0.11 CTIFLCR 0.00 T6M2 0.04 T1M2 0.01 T6M1 -0.04 T6M2 ** -0.11 T6M6 0.37 T6M1 0.26 * T6PHL 0.08 T6M6 0.01 T6PHR 0.09 T6M9 -0.'18 T6TPW -0.06 T6PHL ** -0.11 T1OM2 0.26 T1OM2 *' -0.02 T1OM6 0.38 T10M'l * 0.19 * T1OM9 0.19 T1OM6 o.22 " L1M2 -0.14 T'10M9 -0.07 L1M6 0.24 " TlOPHL * -0.16 L1M9 0.23 TlOPHR 0.14 L5M2 -0.05 T10M11 -o.21 * LsM1 0,14 L1M2 -0.11 L5M9 -0.06 L1M1 0.04 L5M6 0.15 L1M6 0.01 L5PHR 0.09 L1M9 -0.09 15M11 0.1'l L1 PHL 0.09 LsTPW 0.15 L1M1O 11M11 0.09 LsM2 -0.14 LsM1 0.05 L5M6 -0.10 ** LsM9 -0.28 * L5PHL -o.24

Humerus minimal circumference: - females Í Variable - males r Variable 0.14 c7M2 0.13 C7M2 0.22 c7M6 0.14 c7M6 0.01 c7M9 -0.16 CTPHL CTSPL 0.11

*= significant before Bonferroni's correction (p<0'05) **= signif icant after Bonf erroni's correction (p<0'05) Bold: increase (significants only) italic: decrease (signif icants only) 434 F. J. Rühti - Osteometric Variation of the Human Spine 14. Principal comPonent analYsis Males Total Variance ExPlained

Exl Sums Loadi o/ Cumulative % Co Total /o of Variance 19.100 20.538 20.538 32.881 2 11.479 12.343 38.857 ,) 5.558 5.976 43.971 4 4.756 5.1 14 48 5 2 4.6 Extraction Method: Principal Component Analysis.

Component Matrixa

Co

1 2 741 -2.569Ê-02 c3M1 ,737 -.141 TlTPW .690 -..1 18 c3M2 .681 -.138 C3PHR .673 -.399 c7M1 .673 -.119 T1 PHL .655 -.397 T6M2 .648 -.210 T1M1O .643 .425 C3PHL .641 -.347 c3M'11 .632 .315 T6PHR .630 -.508 T1 M2 .621 -.171 CTIFRCA .601 .369 T1 PHR .600 -.226 T1OMg .600 -.244 T6PHL .599 -.427 L5M9 .598 -.124 T6M6 .591 -.355 CTPHL .585 -.211 LSIFRCR .578 .345 T10M10 .573 .453 LSPHR .5Þ | -.294 Tl IFRCA .550 .435 L5IFLCR .543 .140 Tl IFLCA .533 .328 CTPHR .531 -.1 58 C3TPW .524 2.2118-02 T1 OPHR .521 -1.043E-02 L5M6 .513 -.457 T6TPW .512

435 Spine F. J. Ri¡hti - Osteometric Variation of the Hwnan Component

1 2 1 2 -.371 2 .512 -2.884 -03 T1 M9 .345 -7.1698-02 TlOIFLCA .511 .395 T6SPL .143 T1M1 .510 -.255 T6IFLCA .217 8.506E-02 '' -.226 T6M9 .506 -.476 c7M9 .377 -.390 Tl IFRCR .479 .428 c7M6 .389 LSPHL .477 -,315 T6IFRCA 9.026E-02 2.9738-02 -.360 c3M9 .473 1.548E-02 T6M1 .283 -4.212E-02 T1M11 .472 .260 LsM1 .346 L1 PHL .471 -.355 T1OM1 .171 9.2658-02 L1M2 .453 -1 .163E-03 Ll IFRCA .384 .424 TlOPHL .453 -.143 LlTPW -8.837E-02 -1.991E-02 L5TPW .441 -7.968E-02 CTSPL .271 6.763E-02 -4.656E-02 -8.793E-02 T1 M6 .407 -.398 L1 SPL Ll IFRCR .406 .147 L5IFLCA .297 .396 C3IFLCA .384 .379 LSIFRCA .138 .395 TlOTPW .330 -3.407E-O2 T1 SPL .203 .217 -.155 L1M1 .292 -6.O21E-Oz TlOSPL 3.798E-O2 L1M1O .450 .633 Extraction Method: Principal Component Analysis. c7M10 .532 .602 a. 5 components extracted. Ll IFLCR .201 .583 Tl IFLCR .336 .582 T10M11 .424 .561 C3IFRCR .251 .549 c3M10 .268 .545 T6M11 .268 .537 CTIFLCR -6.648E-02 .524 L5M2 .381 -.519 15M11 .470 .508 CTIFRCR 8.430E-02 .497 C3IFRCA .374 .475 C3IFLCR .207 ,472 CTIFLCA .391 .469 TlOIFRCA .356 .469 L1M9 .430 -.464 T1OM6 .411 -.456 L1 PHR .448 -.452 L1M6 .265 -.441 c3M6 .424 -.43s Ll IFLCA .348 .427 c7M11 .328 .404 11M11 .305 .404 LsM1O 6.931E-02 .385 C3SPL .178 -.344 LSSPL -5.713E-02 -.110 T6M1O

F. J. RühIi - Osleometric Variation of the Hurnan Spine 436 Females Total Variance ExPIained

Extraction ums of ared Loadin % Co Total % of Variance Cumulative 26.985 1 25.096 26.985 15 2 11.282 12.131 39.1 45.864 3 6.276 6.749 4 5.140 5.527 51.391 56.731 5 4.967 5.340 Extraction Method: Principal Component Analysis.

Component Matrixa

1 2 7 -.397 c3M1 .354 -.493 c3M6 .274 -.361 c3M9 .548 -7.784F-02 CSPHL .684 -.352 C3PHR .763 -.355 C3M1O .636 .321 c3M11 .647 .203 C3SPL .157 6.055E-02 C3TPW .499 -.251 C3IFLCR .538 .329 C3IFLCA .626 .444 CSIFRCR .504 .510 C3IFRCA .381 .384 c7M2 .808 -.253 c7M1 .724 -.213 c7M6 .547 -.325 c7M9 .372 -.373 CTPHL .503 -.262 CTPHR .551 -.249 c7M10 .634 ,425 c7M11 .447 .333 CTSPL 6.012E-02 .277 CTIFLCR .629 .249 CTIFLCA .454 .626 CTIFRCR .637 .488 CTIFRCA .429 .614 T1M2 .801 -.249 T1M1 .648 -.250 T1M6 .492 -.229 T1M9 .149

437 F. J. Rühli - Osteometric Variation of the Human Spine Component Matrixa

nt

1 2 'l 2 .473 .637 .575 -.253 1 Ll IFRCR .4't6 .245 T1 PHR .676 9:i 4sE-03 Ll IFRCA .562 .s00 T1M1O .664 .400 L5M2 .406 -.193 T1M,I1 .487 .170 LsM1 .448 -.137 T1 SPL .348 -.185 L5M6 .668 -.217 TlTPW .410 -.112 L5M9 .669 -.117 Tl IFLCR .572 .612 L5PHL .751 -5.708E-02 Tl IFLCA .498 .633 LSPHR .817 -.157 Tl IFRCR .623 .428 L5M1O .373 .120 Tl IFRCA .440 .496 15M11 .607 .143 T6M2 .556 -.454 LsSPL 7.559E-02 -.462 T6M1 .346 -.529 LsTPW 5.843E-02 6.868E-02 T6M6 .470 -6.339E-02 L5IFLCR .478 .484 T6M9 .456 -.525 LsIFLCA .165 .539 T6PHL .630 -.246 L5IFRCR .419 .658 T6PHR .673 -.103 L5IFRCA .268 T6M1O .294 .212 Method: Principal Compon ent AnalYsis T6M11 .570 2.183E-03 Extraction extracted. T6SPL .186 2.8378-O2 a. 5 comPonents T6TPW .510 -4.9628-O2 T6IFLCA .144 -.144 T6IFRCA .406 8.234Ê-02 T1OM2 .499 -.319 T1OM1 .216 . -.443 T1OM6 .511 -.235 T1OMg .440 -.482 TlOPHL .588 -9.590E-02 TlOPHR .654 -.248 T10M10 .559 .396 T10M11 .719 -5.2148-O2 TlOSPL .274 -.317 TlOTPW .427 .122 Tl OIFLCA .202 .423 TlOIFRCA .289 .566 L1M2 .507 -.124 L1M1 .488 -.482 L1M6 .529 -.240 L1M9 .556 -.512 L1 PHL .653 -8.866E-04 L1 PHR .581 -.255 L1M1O .754 .231 L1M11 .576 -.187 LlSPL .383 -.328 LlTPW .189 -.406 Ll IFLCR .397

438 F. J. Rühli - Osteometric Variation of the Human Spine

Rühli, F.J., Schultz, M., and Henneberg, M., (2002) Microevolution of the central European human vertebral column since the Neolithic: preliminary osteometric assessment and interpretations. American Journal of Physical Anthropology, suppl. 34, pp. 134-135.

NOTE: This publication is included on page 439 in the print copy of the thesis held in the University of Adelaide Library.

It is also available online to authorised users at:

http://dx.doi.org/10.1002/ajpa.20014