Postcranial Sexual Dimorphism in Pan Troglodytes - UOC THESIS 2012 NOELLE TANKARD

A dissertation submitted in partial fulfilment of the requirement for a
Master of Philosophy in Human Evolutionary Studies
Supervisor: Jay Stock

Word Count: 20,079
Postcranial Sexual Dimorphism

in Pan troglodytes
A detailed consideration of sex differences in a
sample of Pan troglodytes verus from Cameroon
2012
Noelle J. Tankard
University of Cambridge
Department of Archaeology and Anthropology
Division of Biological Anthropology
Pan troglodytes' Postcranial Sexual Dimorphism
Abstract
This study presents information regarding postcranial sexual dimorphism in Pan troglodytes. From a
sample of fifty-six adult chimpanzee specimens of known age and sex (36 female, 20 male), originally from
Cameroon and made available for study at the Powell-Cotton Museum, a comprehensive pattern of
postcranial sex differences is illustrated. This study increases the number of skeletal dimensions with proven
statistically significant sexual dimorphism in the chimpanzee, not only in dimensions not examined in
previous studies, but also with dimensions found insignificant in previous considerations.
Linear measurements of the scapulae, clavicles, vertebrae, humeri, radii, ulnae, ossa coxae, femora,
and tibiae were taken using sliding digital callipers and an osteometric board. Dimensions were chosen to
provide a thorough picture involving size, shape, and robusticity throughout the body, with particular
attention paid to the shoulders and trunk. Statistical analysis of each dimension by sex is reported,
compared to previous work, and discussed. Dimensions are considered as proportions, with ratios and
indices calculated throughout the body and within each element. The ability of various skeletal elements
and anatomical regions to correctly classify sex is addressed with discriminant function analysis. All results
are compared to data from the literature, with both functional and taxonomic implications discussed.
Substantial sex differences in joint surfaces and areas of articulation in the shoulder girdle and
pelvis, long bone lengths, and diaphyseal diameters, were found. The amount of sex difference was in many
instances less than previous studies, but the pattern more pervasive throughout the body. Accuracy in sex
identification of more than 85% is achieved using single skeletal elements, and measurements taken from
the combined elements of the hindlimb classify specimens with 91.5% success. A number of proportions
displayed significant dimorphism and size-adjusted dimensions were able to classify sex with 85.25%
accuracy, clearly demonstrating the non-allometric nature of sexual dimorphism in this population.
Theoretical ramifications include models of palaeoanthropological reconstruction, environmental
and ecological adaptation, and primate intra-specific variability. The results of this study categorically
demonstrate that sexual dimorphism in Pan troglodytes has been under-appreciated in extent if not also in
degree.
2
Table of Contents
Abstract.................................................................................................................................................................................2
Table of Contents................................................................................................................................................................3
Index of Tables......................................................................................................................................................................4
Index of Figures.....................................................................................................................................................................5
Introduction..........................................................................................................................................................................6
Background...........................................................................................................................................................................7
Sexual dimorphism....................................................................................................................................................7
Sexual selection...................................................................................................................................................7
Sexual dimorphism in primates...........................................................................................................................8
Human sexual dimorphism..................................................................................................................................9
Plasticity and development...............................................................................................................................10
Palaeoanthropological implications..................................................................................................................11
Notes on Terminology:............................................................................................................................................11
Quantifying sexual dimorphism.........................................................................................................................12
Chimpanzees...........................................................................................................................................................13
Sex differences in behaviour and sociality.........................................................................................................13
Geographic variation in chimpanzees and subspecies......................................................................................16
Chimpanzee growth...........................................................................................................................................17
Body size: dimorphism and proportions ...........................................................................................................18
Cranial dimorphism ...........................................................................................................................................19
Literature Review: Previous studies of postcranial sexual dimorphism.............................................................................21
Methods..............................................................................................................................................................................31
Sample.....................................................................................................................................................................31
Data collection.........................................................................................................................................................31
Analysis....................................................................................................................................................................32
Results.................................................................................................................................................................................34
Shoulder girdle........................................................................................................................................................43
Pelvis........................................................................................................................................................................48
Forelimb...................................................................................................................................................................51
Hindlimb..................................................................................................................................................................57
Body proportions.....................................................................................................................................................62
Discussion and Conclusions................................................................................................................................................67
Measured values, comparison to previous studies.................................................................................................67
Dimensions of and sex differences in the shoulder girdle.................................................................................67
Dimensions of and sex differences in the pelvis................................................................................................68
Dimensions of and sex differences in long bones.............................................................................................68
Trunk size.................................................................................................................................................................70
Joints........................................................................................................................................................................72
Long bone: lengths and diaphyseal properties.......................................................................................................73
Sex identification.....................................................................................................................................................74
Implications for intra-specific variation...................................................................................................................75
Bibliography........................................................................................................................................................................78
3
Index of Tables
Table 1: Data from previously published studies................................................................................................................29
Table 2: Measurements of the scapula...............................................................................................................................35
Table 3: Measurements of the clavicle ..............................................................................................................................36
Table 4: Measurements of the pelvis..................................................................................................................................37
Table 5: Measurements of the humerus, ulna, and radius.................................................................................................38
Table 6: Measurements of the femur and tibia..................................................................................................................39
Table 7: Limb lengths, ratios, and indices...........................................................................................................................40
Table 8: Sexual dimorphism in proportions........................................................................................................................41
Table 9: Biometric data from field notes, Merfield 1933...................................................................................................42
Table 10: Sex differences in the scapula.............................................................................................................................43
Table 11: Scapular Model: discriminant function analysis.................................................................................................43
Table 12: Sex differences in the clavicle.............................................................................................................................45
Table 13: Clavicular Model: discriminant function analysis................................................................................................45
Table 14: Shoulder Girdle Model: discriminant function analysis......................................................................................46
Table 15: Sex differences in the pelvis................................................................................................................................48
Table 16: Pelvic Model: discriminant function analysis......................................................................................................48
Table 17: Sex differences in the humerus...........................................................................................................................51
Table 18: Humeral Model: discriminant function analysis.................................................................................................52
Table 19: Sex differences in the ulna and radius................................................................................................................54
Table 20: Radial Model: discriminant function analysis.....................................................................................................54
Table 21: Forelimb Model: discriminant function analysis.................................................................................................56
Table 22: Sex differences in the femur...............................................................................................................................57
Table 23: Femoral Model: discriminant function analysis..................................................................................................59
Table 24: Sex differences in the tibia..................................................................................................................................59
Table 25: Tibial Model: discriminant function analysis......................................................................................................59
Table 26: Hindlimb Model: discriminant function analysis................................................................................................61
Table 27: Trunk Model: discriminant function analysis......................................................................................................64
Table 28: Proportional Model: discriminant function analysis...........................................................................................65
4
Index of Figures
Figure 1: Scapula lengths by sex.........................................................................................................................................45
Figure 2: Scapula breadths, by sex......................................................................................................................................45
Figure 3: Glenoid fossa length............................................................................................................................................45
Figure 4: Glenoid fossa breadth..........................................................................................................................................45
Figure 5: Clavicle length, by sex..........................................................................................................................................48
Figure 6: Clavicle mid-shaft diameter (maximum), by sex.................................................................................................48
Figure 7: Clavicle mid-shaft diameter (minimum), by sex..................................................................................................48
Figure 8: Conoid tubercle diameter (anterior-posterior), by sex.......................................................................................48
Figure 9: Clavicle sternal facet height (superior-inferior)...................................................................................................48
Figure 10: Ilium breadth, by sex.........................................................................................................................................49
Figure 11: Iliac crest (minimum) breadth, by sex...............................................................................................................49
Figure 12: Acetabulum height, by sex.................................................................................................................................49
Figure 13: Pubic symphysis breadth, by sex.......................................................................................................................49
Figure 14: Auricular surface length, by sex.........................................................................................................................49
Figure 15: Humerus proximal transverse diameter, by sex...............................................................................................53
Figure 16: Humerus proximal head diameter, by sex.........................................................................................................53
Figure 17: Humerus surgical neck diameter (maximum), by sex .......................................................................................54
Figure 18: Humerus surgical neck diameter (minimum), by sex .......................................................................................54
Figure 19: Humerus mid-shaft diameter (maximum), by sex ............................................................................................54
Figure 20: Humerus distal articular breadth by sex ...........................................................................................................54
Figure 21: Humerus epicondylar breadth, by sex ..............................................................................................................54
Figure 22: Humerus 35% shaft diameter (minimum).........................................................................................................54
Figure 23: Radius height, by sex ........................................................................................................................................56
Figure 24: Radius mid-shaft diameter (maximum), by sex.................................................................................................56
Figure 25: Radius mid-shaft diameter (minimum), by sex.................................................................................................56
Figure 26: Radius head diameter(maximum), by sex.........................................................................................................56
Figure 27: Femur head diameter (maximum), by sex ........................................................................................................59
Figure 28: Femur biomechanical neck length, by sex ........................................................................................................59
Figure 29: Femur distal diameter (anterior-posterior), by sex...........................................................................................59
Figure 30: Femur epicondylar breadth, by sex ..................................................................................................................59
Figure 31: Femur neck length (shortest distance between head and lesser trochanter), by sex .....................................59
Figure 32: Tbia mid-shaft diameter (maximum), by sex ....................................................................................................61
Figure 33: Tibia mid-shaft diameter (minimum), by sex ...................................................................................................61
Figure 34: Tibia plateau breadth, by sex.............................................................................................................................61
Figure 35: “Belly” measurements for a subset of the sample, from 1933 field notes.......................................................63
Figure 36: “Bicep girth” measurements for a subset of the sample, from 1933 field notes.............................................63
Figure 37: “Chest” circumference measurements for a subset of the sample, from 1933 field notes..............................63
Figure 38: “Girth” measurements for a subset of the sample, from 1933 field notes.......................................................65
Figure 39: “Forearm” length measurements for a subset of the sample, from 1933 field notes......................................65
Figure 40: “Hand” height measurements for a subset of the sample, from 1933 field notes...........................................63
Figure 41: “Height” measurements for a subset of the sample, from 1933 field notes....................................................63
Figure 42: “Neck” circumference measurements for a subset of the sample, from 1933 field notes...............................63
Figure 43: “Span” measurements for a subset of the sample, from 1933 field notes.......................................................63
Figure 44: “Hand” height measurements for a subset of the sample, from 1933 field notes...........................................64
Figure 45: “Height” measurements for a subset of the sample, from 1933 field notes....................................................64
Figure 46: “Span” measurements for a subset of the sample, from 1933 field notes.......................................................64
Figure 47: “Neck” circumference measurements for a subset of the sample, from 1933 field notes...............................64
5
Introduction
Chimpanzees, as the living species most closely related to our own, are of great anthropological
interest. They are in an interesting position for scrutiny – it might be more accurate to say that we are in an
interesting position from which to scrutinize them. Our affinity makes study impossible without
comparison; such comparison makes study all the more paramount, but has, throughout history, rendered it
both philosophically and politically volatile. Perhaps it is just that is that which may help explain the irony
that, for a species which has been so richly studied, the sum of our understanding is, today, anything but
rich. In few cases is this more obvious than when it comes to assessing intra-specific variation and sex
differences.
While chimpanzees can be considered, in many ways, to have more sexual dimorphism than
humans, the pattern of sex differences is quite different: there is a far greater difference in the size of male
and female canine teeth, and considerably less difference in their pelvic morphology. As methods of sex
identification used for human skeletal remains are heavily dependent upon the pelvis, they are inadequate
for the identification of sex in chimpanzees. Traditionally, chimpanzee skeletal remains are sexed only by the
canines; the variation in long bones has been considered insufficient to distinguish between the sexes. In
life, chimpanzees show notable dimorphism in body mass. This difference is mainly in girth, as males are
broader than females. A review of the literature, particularly previous morphological studies of various
chimpanzee populations, indicates that sexual dimorphism is greater than has been traditionally
appreciated – and considerably more variable. Skeletal sex differences seem to present in mosaic patterns,
with different populations having different patterns and degrees of sexual dimorphism.
This study is an attempt to provide a more detailed and rich picture of sexual dimorphism in the
chimpanzee postcranial skeleton. Having identified anatomical regions of interest from the literature, I
collected linear measurements from a broad range of skeletal elements. The intention is to further previous
work by increasing the number of anatomical regions considered, including measurements that will allow
consideration of shape as well as size. I hope to provide data that will contribute to understanding of
chimpanzee variation, primate intra-specific variation, and primate sexual dimorphism. I have discussed my
findings with reference to primate evolution allometry and refer to literature regarding plasticity, and
development as related to activity, nutrition, and environment.
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Background
There is a great wealth of anthropological literature regarding primate sexual dimorphism; my goal
here is to provide a brief scope of the trends that have characterized research and to introduce theoretical
and explanatory models, for the purpose of creating a context in which the results of this study can be
discussed. Some information on chimpanzee anatomy, variation, and behaviour is presented with the same
intention.
Sexual dimorphism
Sexual dimorphism is a deceptively simple concept. Common to a wide range of species, from plants
to humans, at the most basic, differentiation in reproductive roles leads to morphological differentiation.
Anisogamy, the separation of the sexes, involves a broad suite of structures, from hormonal pathways to
organ structures. Differential parental investment in offspring and mating strategies continue to modify sex
differences. (Fairbain 1997) Sex differences often take the form of sexual size dimorphism, in which one sex
of a species is larger than the other. Male-biased size dimorphism is the common state for plants. Within
the animal kingdom, it is only in birds and mammals that males predominantly exceed females in size.
(Lindenfors & Tullberg 1998) Despite its rarity for animals, female-biased dimorphism is often referred to as
“reversed sexual size dimorphism” (Ranta 1994, 313). Sexual size dimorphism can vary within taxa as much
as between taxa, but when species are compared within in a clade, sexual size dimorphism is frequently
found to vary allometrically with overall body size. (Gordon 2006b; Ipiña & Durand 2010; Leigh 1992; Leigh
1995; Leuteneggger 1978) This trend has been formalized as “Rensch's Rule”, which states that sexual size
dimorphism increases with size in male-biased cases but decreases with size when females are the larger
(Fairbain 1997, 659). Assessments of Rench's Rule seem to indicate more frequent departures from it in
cases of female-biased sexual size dimorphism. (Abouheif 1997) Allometry, or the “departure from
geometric similarity”, is a description of situations in which differences in proportions are correlated with
changes in the absolute size. (Abouheif 1997)
Sexual selection
Sexual selection, the process by which mate selection and reproductive success exert an
evolutionary pressure on body form, further exaggerates sex differences, resulting in physiological
phenomena that are seemingly far removed from direct reproductive function – the peacock's tail is a
classic example. Secondary sex differences are generally considered to have resulted from sexual selection if
there is reason to believe that they would have increased the bearer's mating success, traits that are either
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preferred by the opposite sex or that give an advantage in intra-sexual competition. Some features, notably
those which serve to display competitive ability, are reasonably simple to identify and postulate as to the
evolutionary pressures which resulted in their development; others are considerably more complex to
identify, much less to speculate with any certainty as to their origin. Furthermore, it is entirely feasible that
any trait, particularly one demonstrating positive allometry, serves multiple functions. (Petrie 1992, 173)
Sexual dimorphism in primates

Primates, including humans, are sexually dimorphic. In most cercopithecoid species, males are
anywhere from thirty to eighty percent larger than females. Of the great apes, gorillas and orangutans are
known for the most extreme body size dimorphism. Humans display only a slight degree of sexual
dimorphism, while that of chimpanzee's is often characterized as moderate. Chimpanzees demonstrate
considerably more dimorphism in canine size than throughout the rest of their body. Compared to humans
and to the Homo genus, chimpanzees have substantial canine dimorphism but compared other primates,
chimpanzees display only minimal canine dimorphism (Plavcan 2001, 48).
Some general patterns in primates are readily apparent from comparing species' size data.
Monogamous primate species generally show little to no sexual dimorphism, with a few exceptions. Species
with high levels of polygyny tend to have higher levels of sexual dimorphism. This correlation is due to a
variety of mechanisms – not all playing equally significant roles in all sexually dimorphic polygynous species
– including intra-sexual competition between males within the group, inter-group conflict as males to
defend group resources (territory and females), and lack of paternal investment in offspring. (Lindenfors &
Tullberg 1998) To conflate socio-economic sex ratio with male-male competition may be premature
(Clutton-Brock 1977). The use of sexual dimorphism as a correlate for mating strategy and social structure is
not without caveats, but it has nonetheless become convention. A competing hypothesis links variation in
primate sexual dimorphism to habitat preference: terrestrial primates tend to be more dimorphic than
arboreal species. Terrestrial species may have a greater need to defend against predators (Leutenegger &
Kelly 1977, in Lindenfors & Tullberg 1998). Arboreal species may be more constrained in size due to their
need to forage on thin tree limbs (Lindenfors & Tullberg 1998). Nocturnal species show less dimorphism
than do diurnal, but there is no apparent difference between folivores and frugivores (Clutton-Brock &
Harvey 1977).
Presuming that sexual dimorphism is entirely a result of changes in male size would presume that
females are, by default, the more “optimal” size for the species in question (Gaulin & Sailer 1985, in
Lindenfors & Tullberg 1998). More recent publications have stressed the contribution of females to body
size dimorphism. Studies continue to demonstrate that sexual dimorphism is not entirely allometric, males
are not simply “scaled-up versions of females”, and sexual dimorphism differs between species in pattern
8
and nature as well as in degree (Plavcan 2011).
Human sexual dimorphism

Human sexual dimorphism is complex; it constitutes not only physiological and anatomical, but the
behavioural, and all three of these can affect each other. In general, all parts of the female skeleton are
smaller and lighter than their respective elements within the male skeleton. Males may average up to 20%
larger in some dimensions, while in others there may be no size dimorphism. Differences in the crania and
pelves are considered the most reliable for sex identification – and often can be distinguished visually by an
observer. Other dimensions, however, can vary by only millimetres. Canine dimorphism is considerably less
dramatic in humans than it is in other apes, and the degree of sexual dimorphism in crown sizes varies
between populations. (White & Folkens 2005)
There are a number of accepted methods for identifying the sex of human skeletal remains. Results
from long bones often result in 80 – 90% accuracy (Rogers (1999). Intra-specific variation in human sexual
dimorphism is so great that it is generally considered unadvisable to attempt to identify the sex of a skeletal
specimen without reference to its population of origin; conversely, when analysing large samples of skeletal
specimens, the magnitude of sex differences can be used as a criterion for establishing the sample’s origin.
Perhaps most intriguingly, while the degree of sexual dimorphism varies between populations, it “does not
seem to differ on the average from region to region” (Frayer & Wolpoff 1985, 430). There is evidence for
secular trends in human and hominid sexual dimorphism, with significant decrease in sexual dimorphism
even between European Upper Palaeolithic groups to Mesolithic and Neolithic populations of similar
regions. Frayer and Wolpoff (1985) have suggested that nutritional changes can explain much of the
temporal fluctuation in human sexual dimorphism – particularly those that coincide with changes in dietary
strategy, as populations shifted from hunting and gathering to agriculture and horticulture.
A quick consideration of human sexual dimorphism reveals the multiplicity of ways in which it is
expressed, and the complexity of the phenomenon. While human body mass dimorphism is relatively
meagre as compared to other species, the slight differences in cranial-facial features, muscle mass, amount
of subcutaneous fat, growth rates, and disease vulnerability that characterize the sexes are only a portion of
that which differentiates them. Furthermore, that there is a great deal of variation between populations in
the degree and nature of sexual dimorphism is another facet which, while taken for granted in our intrinsic
understanding of our own species, is something that is often under-appreciated when considering other
species.
While the possibility that reduced sexual dimorphism in chimpanzees and humans is a primitive
trait, a characteristic of their last common ancestor, the simplest interpretation of the data is that the
reduction of dimorphism in the chimpanzee and human lineages is a homoplasy. Sexual dimorphism seems
9
to drop considerably with Homo erectus, and again with Neanderthals. Postcranial specimens of Homo
habilis seem to demonstrate similar dimorphism as Australopithecus afarensis. The limited nature of Homo
erectus postcranial remains renders any conclusions premature; however, data suggests that body size
dimorphism is reduced (Frayer & Wolpoff 1985, 456)
Plasticity and development

Most sex-linked traits are polygenetic and become fully apparent at puberty (Frayer and Wolpoff
1983; Haqq 1988). The plastic nature of such traits complicates any attempt to draw direct evolutionary
pressures, as they can vary in expression and development, responding to environmental cues and changing
on a temporal scale, in generation as opposed to millennia, that would be inconceivable for other physical
structures. Both genetic and non-genetic (including epigenetic and environmental stimuli) influences must
be considered when attempting to theorize causal mechanism.
Diet and nutrition play a major role in growth. Males are more “susceptible to fluctuations in
nutritional quality and show greater impairment in long bone growth”; that females are less affected and
“more stable under the same food deficits” is, presumably, a result of reproductive necessity (Frayer 1985,
431). The female body, designed to grow to a smaller body size and, both post-pubescent a nd throughout
life, characterized by a greater amount of subcutaneous fat. (Frayer & Wolpoff 1985, 432) If differential
dietary deficiencies are chronic, it would be expected that selection would reduce body size as related to
energetic efficiency; it would also be expected that males would be under the most intense selection.
Nutritional stress is thought to decrease sex differences in humans, while populations with
improved diets seem to have increased sexual dimorphism. There are a number of behavioural factors that
complicate anthropological examinations of the phenomenon, especially when making cross-cultural
comparisons. Many of the contemporary societies in which dietary deficiencies and the resulting growth
problems, i.e. stunting, have been the most thoroughly examined are societies with strong patriarchal
structures, in which female children suffer from disproportionate allocation of resources. (Grantham &
McGregor 2007; Ghosh 2009; Pramod-Singh 2009) This no doubt distorts the effects. However, when
looking at descendent populations the effects of dietary changes can be analysed. African “Bushman”
groups that adopt “Neolithic” subsistence patterns show a marked increase in male stature. Cultural
practices confound the issue, making it unclear the disproportionate increase in male stature in these cases
resulting from the nutritional surfeit relieving the male deficiency, or a differential sex access to the new
agricultural foods, throwing females into a depressed nutritional status. (Frayer & Wolpoff 1985, Wolfe &
Gray 1982) Some generalizations can, however, be drawn; that these generalizations may have cross-special
implications is a theoretical stretch that, nonetheless, merits consideration.
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Palaeoanthropological implications
A large portion of palaeoanthropological debate and evolutionary theory is concerned with
physiological changes that characterize the hominin lineage. Of these, the increase in cranial capacity,
adoption of obligate bipedalism, and reduction in tooth size have received the most attention (Plavcan
2001). Some authors, however, have suggested that the lack of sexual dimorphism, interpreted as a
continued decrease throughout the lineage, may be as critical to defining hominins as an y of the others.
(Frayer & Wolpoff 1985, 451)
The study of sexual dimorphism in the fossil record has been directed, for reasons of obvious
imminence, in distinguishing sex differences from taxonomic differences. Ascertaining how many species
are represented by multiple specimens, or even multiple sites, is an ongoing and often contentious process.
The matter of how much variation can be encompassed within a hominid species is itself an open question.
When the size or shape of a number of specimens presents with an apparently bimodal distribution, it is
not an unreasonable assumption that a sexually dimorphic population is more likely than a population
consisting of multiple clades – however, such hypotheses require further consideration. Whether the size
variation in australopithecine specimens is explained by a high degree of sexual dimorphism or necessitates
a re-evaluation of the number of species which requires an understanding of intra-specific and intra-sexual
variation which, arguably, we do not yet have.
Notes on Terminology:
Throughout the paper, I use the terms “sexual dimorphism” and “sex differences” interchangeably
with “sexual size dimorphism”. That sexual dimorphism in chimpanzees and primates takes the form of
sexual size difference is an obvious starting point; the majority of the sex differences with which this paper
is concerned take the form of size difference, in various skeletal elements and measurable dimensions. Sex
difference is a broad term that encompasses any and all physical differences between a species' sexes, and
is often broken down into two categories. “Primary” sex differences, those directly related to mating and
reproduction (I.e., pelvic morphology necessary for obstetric needs), and “secondary” sex differences, or
any sex-linked trait that does not serve an immediate functional purpose in reproductive behaviour.
Secondary sex differences are primarily discussed as products of sexual selection. In mammals, pelvic
morphology is the most obvious primary sex difference. In primates, the most readily apparent of
secondary sex differences are body size and canine dimorphism. The term “sexual dimorphism” has become
entrenched in sexual selection, as a short-hand reference for such traits. Both body size and canine size
have long been worked into theoretical models which posit their roles in intra-sexual competition (Plavcan
1997); it has even become axiomatic in basic physical anthropology texts that larger canines give males a
11
competitive advantage in fighting for mates, while larger body size is related to a species' social structure
and mating strategy. These generalizations and evolutionary models are eminently useful in inter-specific
comparisons, but they are considerably less so when addressing intra-specific variation. The correlations
between body size and mating strategy are considerably less powerful the more closely-related the taxa
under consideration are to each other, and there is no concrete evidence yet that sexual selection has any
power to explain intra-specific primate variation.
Creating a binary distinction between primary and secondary sex differences reinforces the
perception that all secondary sex differences are related to sexual selection and, therefore, intra-sex
competition. The distinction is not only irrelevant to the purposes of this paper – in which I am attempting
only to detail a thorough description of postcranial sexual dimorphism in a single species – but is perhaps
theoretically ill-advised. It distracts from the truth that it is the same processes of development,
complicated pathways by which androgens interact and by which both primary and secondary sex
differences are created. It depreciates the fact that sex differences not only characterize mature individuals,
but influence behaviour both pre- and post-puberty and, given the growing appreciation of the impact that
development and life history factors have upon the skeletal form, it seems all the more prudent to frame
the discussion in the broadest of terms. Furthermore, the sexually dimorphic postcranial traits which
constitute the focus of this paper have no clear explanatory models and no need to be termed as products
of sexual selection or of reproductive function.
Quantifying sexual dimorphism

There are a number of commonly used formulas to calculate the degree of sexual dimorphism
metric features. The simplest – dividing the mean of one sex by another to create a ratio – has long been
the favoured method in primate studies. Several decades of literature argue that residuals from regression
models are preferable. Ranta (1994) criticizes the use of ratios in assessing sexual dimorphism because they
mask the effects of body size, and sexual dimorphism often increases as body size increases. Plavcan (2011)
echoes these concerns and stresses that more sophisticated statistical analyses reveal considerably more
data, including differences in variances between samples. In this study, as it is working within a single
population, I am simply using basic ratio calculation (of females as a percentage of males) and coefficients
of variation; the former for its power to fully address dimorphism within the sample of this study, and the
latter to ease immediate comparison to previous publications.
12
Chimpanzees
Sex differences in behaviour and sociality
As a species, chimpanzees are both highly independent and highly sociable; they spend large
amounts of time alone, autonomous, relatively private – and yet their interactions and relationships play a
significant role in their lives. “Freedom in choice of companions, travel route, activity, and so on, plays an
important role in reducing stress, particularly for males. At the same time, the constant separation and
reunions impose unusual demands on social intelligence and awareness and exert strong pressures for
flexibility of behaviour (Goodall 1986, 167)”. Chimpanzees have a so-called fission-fusion social structure,
with members of a community breaking into variously composed small groups to travel, feed, and sleep; the
most common are all-male parties, family units (a female with her young offspring, possibly grown
daughters), nursery units (two or more females with their young), courtship parties (a female in oestrous
with a single male), sexual parties (female in oestrous with multiple males, possibly other females), mixed
parties, and larger gatherings.
Males could be considered more sociable than females in the frequency with which they met non-
family individuals, the number of regular associates they keep, and the duration of their associations – they
join more gatherings and stay longer. There was variation in the level of individuals' sociability, but in
general, males spent more days as a member of a party, while females had more days with family members.
Females were more sociable not only when in oestrous themselves, but when their grown daughters were
cycling as well. As males lost status, or dropped in relative dominance, they often became less sociable.
Goodall et al (1986) found that the all male travel parties consisted of, on average, 4 to 5 individuals, while
females were on average in groups of 1.6. Any male is likely to encounter all other males of the community
several times a week, while some females may only see each other a few times a year. Females may migrate
(permanently or temporarily, some move continually back and forth) between communities. When food
was scarce, chimps broke into smaller groups. The availability of females in oestrous was the most
significant factor in the overall community patterning from year to year.
The social lives of juveniles, of either sex, are primarily determined by their mothers' lifestyle. Her
personality determines how often and with whom the juveniles have the opportunity to socialize, but the
result of any aggressive encounter between juveniles is directly determined by their mothers' respective
ranks, while the presence, or lack of thereof, older siblings greatly affects the day to day life of a juvenile.
Males under 8 years are very eager to join other parties – they will beg their mothers, crying and
whimpering if they refuse – particularly if those groups contained there are juveniles of a similar age,
females in oestrous, or sociable adult males. Around 9 years of age, males begin spending less time with
their mother and more time with non-family members; at late adolescence they become somewhat more
13
“peripheral”, rising throughout their adult life as they build relationships with other males. Juvenile females
are somewhat less gregarious than males, less eager to join other groups and less likely to coerce their
mothers to stay with a group; by late adolescence, 10 years and older, they become intensely social,
frequently leaving their mothers to travel with adult males; after the birth of their first child, they spend
more time on their own, or with only their offspring for company. Infant mortality is high for both sexes –
although males are at a greater risk of mortality as adolescents and young adults. Those that survive tend to
live to an older age.
Grooming is the prime example of affiliative behaviour. Males spend more time involved in
grooming sessions with non-family members than do females; they do groom younger and lower ranking
individuals – although less frequently and for shorter intervals than higher-ranking individuals. Grooming is
both a reflection of and a means of establishing alliance relationships; the frequency of grooming sessions
between two males may increase when the relationship is strained. A female may spend more time
engaged in grooming sessions, but she does so with a smaller number of individuals – primarily her family
members and older males; rarely do they groom unrelated females, who even more rarely reciprocate.
Younger individuals more often initiate grooming, but grooming which is not reciprocated ends more
quickly. (Goodall et al 1986)
There are obvious sex differences in aggressive behaviour, both hunting and agonistic. Most
aggressive behaviour consists only of threats; gestures and postures, vocalizations, and displays resolve the
majority of conflicts. Actual fights, when they do occur, are generally brief – the only fights observed by
Goodall and researchers at Gombe which lasted longer than five minutes, were intra-community, with
several members of a community attacking and brutally injuring a foreign individual. Intra-community
aggression tends to only involve males, as females move between groups with relative impunity. “Agonistic”
behaviour, a term coined by Scott and Fredericson (1951) encompasses all intra-specific competition (I.e.
threat, attack, defense, escape, and appeasement), is equally common to males and females, although the
nature and type of attacks differ by sex. When males attack, they often leap onto the back of their
vadversaries and stamp with their feet; females tend to grapple, appearing to wrestle, rolling back and forth
with their victims. Other individuals often become involved in group fights– allies coming to the aid of one
party, or a higher-ranking individual separating those involved. Males attack during reunions, meat-eating,
sexual encounters, and social excitement (many of these attacks no doubt related to establishing,
confirming, or challenging the dominance hierarchy); 82% of their aggression and 88% of attacks are
directed against other adult males. Females, however, attack only in context of feeding and protection (of
offspring); 33% of their aggression is directed at adult males (attacking them only to protect or save their
offspring), 33% at adult females, 21% at adolescent females.
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Chimpanzees primarily meet their nutritional needs through foraging in trees. Males have been
observed to consume more meat than do females. When chimpanzees hunt as a group – the most detailed
accounts being of predatory attacks on baboons, colobus monkeys, and bush pigs – prey is generally killed
by a bite to the head, blunt trauma from being flailed against the ground, or dismemberment. There is
intense aggression surrounding kills, between all members of the group; many will attempt to rip off
portions of a carcass, others will beg. Carcasses are often stolen outright from the individual who caught
them, or at least have large portions immediately torn apart. While males are known for hunting more
frequently and more elaborately – in groups with co-operative strategies – females do hunt as well, both on
their own and with others. When in mixed company, even if successful at catching prey, they are unlikely to
be successful at holding onto their kills. With small prey, a preference for brain tissue and blood is clear, as
these are always consumed first; with larger prey, for which chimpanzees' teeth are not sufficient to kill, the
viscera is eaten first. Chunks of meat are torn from the body and chewed along with wads of leaves – these
inedible “wadges” are discarded, although frequently picked up by other individuals who were unable to
obtain any meat themselves. Large bones are cracked open and the marrow extracted. (Goodall 1986)
Chimpanzee locomotion is primarily quadrupedal, as they travel on the ground between sites. They
feed and sleep in trees; locomotion during feeding is also primarily quadrupedal, but it includes more
bipedal and arm-swinging behaviour. Doran (1993) undertook a study of chimpanzees at Tai Forest, Cote
d'Ivoire to determine sex differences in locomotion, posture, and substrate use – results included found
locomotor and positional differences consistent with body size differences. Comparisons of locomotor
strategies and body size have, for the most part, addressed cross-species differences: in such, Fleagle and
Muttermier (1980) and Crompton (1984) found that, across seven sympatriatic species of monkeys and two
galago species, larger animals leap less, climb more often, use larger substrates, and engage in more
frequent suspensory behaviour, In Doran's study, 430 hours of observations were taken over seven months;
focal animals, from all age and sex classes, were followed over the course of entire days with instantaneous
sampling and locomotor behaviour sampled. Three hypotheses were considered: whether the larger males
leapt less, climbed more, and used suspensory behaviours more frequently than did smaller females,
whether there were apparent sex differences in height preference and substrate use, and whether
positional activity differed between the sexes as related to social and maintenance activities. The site had
been chosen because, as a lowland rainforest, it was thought that arboreal travel would be more important
than at other more terrestrial sites; however, this did not seem to be the case. Travel between feeding and
resting sites occurs on the ground. Arboreal travel is mainly vertical, with only occasional horizontal
movement, from a large tree to one adjacent, smaller tree more suitable for descent – in line with other
published accounts (Kortlandt 1962, 1975; Goodall 1965, 1968; Reynolds and Reynolds 1965; Hunt 1992).
15
Doran (1993) did not find sex differences in overall travel (terrestrial and arboreal together), or in
the frequency of behaviours in feeding or travel locomotion. Terrestrial locomotion was 84% of all sampled
locomotion and showed no significant sex difference; the remaining arboreal locomotion, however, did
differ by sex – but only during feeding, not travel. Adult males used significantly less quadrupedalism
(particularly palmigrade), engaged in more climbing, tree-swaying, and bipedalism than did females, and
were slightly more suspensory. Females had a slight tendency to use more quadrupedalism and less
climbing, scrambling, and aided quadrupedalism; this would be explained by the fact that females, being
smaller, encountered more branches capable of supporting their weight and had less need to recourse to
other behaviours. Males and females do not differ in frequency of substrates used – but, in how these
substrates, particularly branches, are used. Males and females differed in height preference. Suspensory
behaviour itself had no sex differences. Females significantly spend more time resting above ground than do
males. In addition, there were sex differences in postural behaviour, especially during resting – males lie
more and sit less frequently than do females. Sex differences in mean party size and the presence of
predators might contribute to females' tendency to spend resting time above ground; female group size is
generally smaller, and all females observed in this study had infants or juveniles. Inter-site difference might
be related to body size difference, as it is likely that Tai chimps are larger than Mahale or Gombe.
Geographic variation in chimpanzees and subspecies

There are three commonly recognized subspecies of chimpanzees: Pan troglodytes troglodytes (the
“common” chimpanzee of Central Africa), Pan troglodytes schweinfurthii (the East African chimpanzee,
perhaps the best known subspecies due to the long-running studies at Gombe and Mahale National Parks in
Tanzania), and Pan troglodytes verus (the West African chimpanzee, studied at Tai National Park in the Cote
d'Ivoire, also known in Guinea). A fourth subspecies, Pan troglodytes vellerosus, has been suggested based
on genetic evidence. Gonder et al (1997, 2006) have published the results of multiple analyses of
haplotypes from the first hyper-variable region of mitochondrial DNA, sampled from chimpanzees in Nigeria
and Cameroon which lead them to conclude that a “major phylogenetic break between lineages occurs near
the Sanaga River in central Cameroon” (2006, 1103). The existence of Pan troglodytes vellerosus as a
distinct clade is yet to be accepted without hesitation by the primatological community at large (Becquet et
al 2007), but there is already contention regarding the nomenclature. Oates et al (2009) argue that the
original type specimen of Pan troglodytes vellerosus, collected in 1862 by Richard Burton, was actually from
Gabon. They suggest that an available name for a population in the Nigeria-Cameroon region, due to
historical contingencies, would be Pan troglodytes ellioti (2009, 80).
Taxonomic analyses of chimpanzees have a long and varied history, but can be traced back to the
work of Scharwz (1932), in which he collapsed a number of historical taxon names to give us those
16
recognized today (Jabbour 2008, 23). Scharwz built on the work of Allen (1925), who gave a detailed
description of the variation in the skin and skull of chimpanzees from two sites in the Upper Congo. Early
attempts at classification were primarily based on external features, particularly skin and hair colouration.
Advancements in genetics and the recognition of bonobos as a separate species have resulted in
contemporary research, even studies not explicitly designed to explore it, to be nonetheless coloured by an
interest in distinguishing Pan paniscus from Pan troglodytes (Coolidge 1982). Groves (1989) undertook a
revision of great ape systematics, compiling a list of morphological and behavioural diagnostics of
chimpanzee subspecies; his work included few postcranial measurements and no single cranial feature for
all subspecies (Jabbour 2008, 24).
Studies based on cranial features and dentition have had varied results when it comes to evaluating
the relationship of the subspecies.(Jabbour 2008) The existence of at least the three main subspecies are
generally supported, as is the relatively greater separation of Pan troglodytes verus from Pan troglodytes
troglodytes and the affinities of the latter with with Pan troglodytes vellerosus. However, there has been
frequent indication of further variation on a population level, leading some authors to simply discuss
chimpanzee morphological variation in terms of population (Pilbrow 2003; Uchida 1993). Genetic analyses
suggest that Pan troglodytes troglodytes and Pan troglodytes schweinfurthii are more closely related to
each other than either is to Pan troglodytes verus, which diverged at an earlier point in time (Becquet 2007;
Jabbour 2008). Genetics have strengthened the assumption that the chimpanzee subspecies have little
admixture (Becquet 2007; Jabbour 2008; Morin et al 1994; Gagneaux et al 1999).
Studies of postcranial morphology are less clear. Groves (2001) recognized four subspecies, but
provided data only to distinguish Pan troglodytes troglodytes, with relatively long limbs to skull size, from
the “shorter-limbed” Pan troglodytes verus and Pan troglodytes schweinfurthii. Jungers and Susman (1984),
in a study of allometry intended to distinguish Pan paniscus, found that “Pan troglodytes schweinfurthii had
narrower scapula and radial head diameter than male Pan troglodytes troglodytes, female Pan troglodytes
scwhweinfurthii had a narrower distal humerus than female Pan troglodytes troglodytes, and subtle
differences in the growth patterns of the hindlimb and forelimb were revealed” (Jabbour 2008, 27).
Morbeck and Zihlman (1989) quite thoroughly demonstrated geographic variation in Pan troglodytes,
although their results differentiated Gombe Pan troglodytes schweinfurthii from others of the same
subspecies, they did little to clearly advance subspecies diagnostics. Carlson (2005) compared cross-
sectional properties, looking at principle areas of movement in humeral and femoral diaphyses, wth
significant differences distinguishing female Pan troglodytes verus from other subspecies.
Chimpanzee growth
The growth trajectories of male and female chimpanzees differ substantially. Pusey et al (2005)
17
found that body mass dimorphism was minimal to non-existent throughout the first 10 years of life, during
which all juveniles showed the same “more or less exponential” growth. Males continued the same
exponential growth until reaching 12-13 years of age, peaking in mass around age 25 and declining slightly
thereafter. Goodall (1986) reported that the males of Gombe National Park displayed an acceleration in
weight gain between 9-10 years of age, with adult weight reached by 16-17 years of age; there was a
“noticeable” increase in scrotum size at 9-10 years, with full size reached at 12-13 years, when males
become functionally fertile. Females at Gombe reached full adult weight later, at 19 years of age, but
became fertile younger: sexual swellings came at 7-8 years of age, increasing to full-sized swellings at 10-11
years, the first true oestrus, with first menarche approximately six months later. The largest amount of body
mass dimorphism was evident at 13 years of age. Intriguingly, Goodall (1986) notes that Gombe
chimpanzees matured later than captive chimps, explaining the phenomenon by reference to the
differences in body weight. (Pusey 1977; McGinnis 1973)
It seems likely that the pattern of growth is strongly correlated to sexual dimorphism. Martin et al
(1994) suggested that the sort of dimorphism in which females demonstrate reduced body size without
reduced cranial capacity or tooth size – the pattern seen in chimpanzees – implies an extended growth in
males, as brain size and molar tooth size reach adult size earlier in the growth pattern. Leigh and Shea
(1995) compared growth, body mass dimorphism, and behavioural ecology in living hominoids. Their study
described the ontogeny of orangutans (males seemed to grow steadily and consistently), gorillas and
bonobos (females mature early while males continue to grow), and chimpanzees (males have an increased
rate of growth). They further suggested that selection favoured rapid female maturation and an earlier
reproductive start for gorillas and bonobos, species characterised by scramble competition, as opposed to
chimpanzees, a species characterised by female contest competition, a situation in which prolonged growth
minimizes juvenile risks. (Plavcan 2001, 45) Plavcan and van Schaik (1995) attribute the lack of reduced
canine size in females to sexual selection acting on female-female contest competition and a need for
“enhanced female weaponry”. That these two theories hypothesize the same causal factor (female contest
competition) to produce the same effect (increased female canine size) through different mechanisms is
telling of complications involved in evolutionary models.
Body size: dimorphism and proportions

Uehara and Nishida (1987) reported an an average male body mass of 42 kg for males (N = 6) and
35.2 kg for females (N = 8) for Pan troglodytes schweinfurthii of Mahale National Park, Tanzania. The
weights were taken over a seven-year study by luring the chimpanzees up a rope hung from a spring
balance. Weight fluctuated seasonally, consistently higher in the wet seasons than the dry. Data sets from
other populations were similar, with Pan troglodytes schweinfurthii of eastern Zaire averaging 42.8 kg for
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males and 34.3 kg for females (Rahm 1967, in Uehara and Nishida 1987, 319) and Pan troglodytes
schweinfurthii of Gombe National Park averaging 39.5 kg for males and 29.8 kg for females (Wrangham and
Smuts 1980, in Uehara and Nishida 1987, 319). The 83.8% dimorphism in body mass at Mahale was
comparable to the 75.4% at Gombe and 80.1% in eastern Zaire. Differences between the male means by
site were not statistically significant; the female means were not similar, as the females of Gombe were
significantly lower than those of Mahale and Zaire.
Pusey et al (2005) undertook a study of body mass in wild chimps, looking at ecological and social
factors affecting variation and fluctuations in body mass. Pan troglodytes schweinfurthii at Gombe had
median male body mass of 39 kg (N = 31) and a median female body mass of 31.3 kg (N = 26), over the
course of 33 years of study. Body masses fluctuated between years and between seasons; all individuals
were heavier when the range size was large and the population density low, and heavier in the wet season
than the dry. Higher-ranking individuals of both sexes tended to maintain more stable body mass than
lower-ranking individuals, but it was only for females that dominance rank was significantly correlated with
body mass. It was unclear as to whether this relationship was due to higher-ranking females having priority
access to food, or if their higher body mass was a factor in their achieving or maintaining body mass (Pusey
et al 2005, 26). Females gained approximately 15% of their body mass during pregnancy; no significant
change during lactation appeared in the dataset, but as many measurements during lactation were taken of
the mother and infant combined, with a rough estimation mass of offspring removed in calculation, the
data may have been too crude to accurately determine mass (Pusey et al 2005, 24).
That the chimpanzee group at Gombe experiences a higher population density from loss of habitat
has been suggested to explain the differences between the chimpanzees of that location from others of the
same subspecies. Morbeck (1999) found evidence for a secular change in chimpanzee size, with individuals
born before significant human disturbance in the area apparently larger (in Pusey et al 2005, 24).
Cranial dimorphism
Skeletons of adult chimpanzees are most reliably sexed by their canines. Males' are larger and
longer, and their maxillary bones broader., to incorporate the larger alveoli (Dierbach 1986, 45). Medial and
lateral incisors are smaller in females (James 1960). Post-canine dentition displays a small amount of
dimorphism, deemed insufficient for sex identification (Zieglansberger & Brandt 1974; Almquist 1974;
Wood 1991). The mandible is generally more “strongly developed” in males, but not measurably so as to
significantly differentiate sex, apart from the lower canines (Dierbach 1986, 45). The larger masticatory
process of male chimpanzees forces the zygomatic arch to project more laterally (Dierback 1986, 45). In the
neurocranium, the crista occipitalis is more strongly pronounced in the male; the weak development of the
crest results in the female occiptal being more rounded (Dierbck 1986, 46). Some male chimpanzees have a
19
quasi-sagittal crest, from the convergence of the lineae temporales, but none was seen in the sample
studied by Dierbach (1986).
Dierbach (1986) examined a collection of 277 Pan troglodytes verus skulls (100 male, 129 female). A
large number of landmarks on the canines, maxillas, mandibles, and occipitals were measured and
analysed. In all dimensions examined by Dierbach the mean values for males exceeded the female means,
which usually exceeded the male minimum – in a few instances, the female maxima reached or even
exceeded the male. The skulls of males averaged 3.7% larger than female, but the amount of overlap
between the sexes was considerable. Dierbach concluded that the degree of sexual dimorphism in Pan
troglodytes verus is less than they had expected – meaning, less than was known in previous studies of
other subsepcies.
Wood et al (1991) compared cranial dimorphism in Homo sapiens, Gorilla gorilla, and Pongo
pygmaeus to a collection of Pan troglodytes verus from the Powell-Cotton Museum – the same collection
which is measured in this study. A total of 79 variables were analysed; nearly all of these demonstrated
statistically significant sex differences in gorillas and in humans and orangutans, both at 80%, although with
different dimensions. In chimpanzees, only just over 50% were statistically significant.
Fenart and Deblock (1973) looked only at the Pan troglodytes collection of the Belgian Royal
Museum of Central Africa, 19 male and 23 female skulls. Of a set of 342 measurements 98 of these
dimensions showed highly significant sex differences (p-values < 0.01), with an additional 37 dimensions
significant (p-values <0.05). Generally speaking, the dimensions with sex differences are related to the
overall larger size of the male cranium. In addition, the naso-frontal area is more advanced in males, who
have larger facial mass. Mandibles are larger in length, height of the ascending branch, and posterior and
anterior widths.
The comparison of the chimpanzees of Tai and Gombe undertaken by Zihlman et al (2008) revealed
that the degree of cranial dimorphism is not standard across all chimpanzee populations and subspecies.
Cranial capacity differed significantly at Tai but not at Gombe; males were similar across the sites, but the
females at Tai were significantly smaller than those at Gombe. The lack of dimorphism in cranial capacity at
Gombe did not preclude significance of sex differences in other cranial-facial dimensions (despite evidence
that cranial-facial differences result from variation in skull size): bizygomtic breadth, mandibular length, and
facial height. Nonetheless, of the cranial-facial dimensions considered in the study, a larger number were
significantly dimorphic at Tai (including palatal length and interorbital breadth, in addition to bizygomatic
breadth, mandibular length, and facial height) than at Gombe. Between the sites, palate length, mandibular
height and length, and interorbital breadth differed significantly – implying a greater plasticity in these
regions.
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Literature Review: Previous studies of postcranial sexual

dimorphism
While there is a long history of studying sexual dimorphism in primates, within the context of
comparison to humans, there have been relatively few studies focusing solely on extant ape species, much
less chimpanzees. Adolph Schultz devoted his career to documenting primate anatomy. The Schultz
collection of specimens, not to mention his work quantifying the variation therein, are still among the most
thorough. In a 1949 paper on the sexual differentiation of primate pelves, Schultz discussed earlier work on
the topic – namely, studies by a German researcher published in 1911 and 1914. While van de Broek's
initially concluded that “practically all the pelvic characters which tend to separate the sexes in man show
also more or less marked sex differentiation in the non-human primates” (Schultz 1949, 402), Schultz
argued that the data reported in van den Broek's second paper, which he characterized as a more thorough
and detailed assessment of the matter, did not support the initial claims. The analysis had been based on a
very small sample which included juveniles. While subsequent work has been equivocal when it comes to
the degree of sexual dimorphism in primates, it has been incontrovertibly demonstrated that there is a
great deal of variation within the primate family. Inasmuch as a theme could be claimed, research continues
to emphasize that the mosaic nature of the differences between monkeys and apes, not to mention
between different species of apes – which is to say, non-human primates do not display the same
charactersistic features of sexual dimorphism, in the pelvis or elsewhere, as do humans.
William Straus (1929) published a series of papers on the phylogeny of the ilium; in one focusing on
the primate ilia, he compared humans, “anthropoid apes”, Old World Monkeys, New World Monkeys, and
Lemuroidea. The comparison was broad, but found consistent sex differences for anthropoid apes and
macaques, in that the portion of the lilium lying above the sacro-iliac joint is proportionately larger in males
than females (Straus 1929, 403). Schultz found similar results for proboscis monkeys, gibbons, orangutans,
and chimpanzees, in a variety of additional pelvic indices, including the breadth of the pelvic inlet in relation
to trunk height, and the maximum pelvic breadth (1949, 402). The ischium-pubis index, introduced by
Schultz (1930), was used by various researchers thereafter. In 1942, Washburn compared skeletal
proportions of langurs and macaques, finding that the pubic bones of females were longer than those of
males. In order to compare across species, he used Schultz's ishium-pubis index and found that, while the
ischium varied in length proportionately to body size, the pubis was disproportionately longer in females.
(1948, 199). A subsequent paper in 1948, which aimed at “present[ing] a method of sexing skeletons which
works in a greater percentage of cases” expanded the analysis to 300 Homo sapiens specimens of multiple
21
races, using the same set of measurements that had been originally modelled after Schultz (1930). While
the ischium was both relatively and absolutely longer in females, the degree of overlap between ranges
differed between human races; overall, the mean difference (between male and female sizes) in man was
less than the mean difference in macaques (1948, 201).
Schultz's 1949 study was expressely designed to extend Washburn's work, to “gain more
information on the ischium-pubis index in a larger variety of primate species and to compare both these
pelvic indices in the same series of specimens whenever possible” (403). A wide range of species were
considered (black-handed spider monkeys, rhesus monkeys, proboscis monkeys, white-handed gibbons,
silvery gibbons, orangutans, chimpanzees, gorillas, and humans); the ischium-pubis index was calculated
for 479 specimens, and the relative pelvic inlet breadth for 485 specimens (404). Of the 88 Pan troglodytes
specimens (21 female, 20 male) (405) considered for analysis, an additional 9 sub-adult specimens were
measured; several (precise number not specified in the paper) had died in captivity rather than having been
wild-shot – Schultz was not concerned that this would affect the results, as “these do not include any pelves
suspected of having been affected by any pathological process” (404). Analysis showed that, generally,
averages for adult females surpassed those of males in both the minimum and maximum values; they
showed considerably more variation, for all indices in all monkey, ape, and human species. Furthermore,
there was very little difference in sub-adults male and females, so it appeared that pelvic dimorphism
resulted from growth that was more intense in females than in males (407).
The results for the chimpanzees, however, stood out significantly from all other species in several
ways: the chimpanzees showed no sex difference in pubis length and ischium length, which normally
averages longer in males than females, was smaller for the male sub-adult chimpanzees. (406) Schultz
concluded that African apes demonstrated considerably fewer sex differences in pelvic shape than did other
primate species – so much so that he declared that “in chimpanzees and gorillas, therefore, the relative
length of the pubic bone is practically useless for diagnosing sex. In orang-utan and gibbon the sex
differences in the ischium-pubis index are clearly more marked, though much less pronounced than in
monkeys and man (411)”, he found that the relative pelvic inlet breadth had comparatively more sexual
dimorphism in the African apes, macaques, and humans (as opposed to the smaller sex differences in spider
monkeys and gibbons). However, the overlapping between the ranges of male and female averages
rendered the differences in pelvic inlet breadth unsatisfactory for determining the sex of a specimen (411).
Schultz turned to functional considerations, namely the size of each species' newborns relative to
the female pelvis, for the majority of his discussion regarding primate variation. He noted, however, that in
this relationship humans seemed to occupy “an intermediate position” between monkeys and the other
great apes (416); he characterized the “anthropoid apes” as an exception to the rule that pelvic size
22
differences are primarily a result of infants head size (418). If the apes are an exception to other primates in
this manner, chimpanzees seem to be then an exception to the exception. In his limited discussion focusing
directly on chimpanzees, he did allege that it was highly significant that chimpanzees showed smaller
secondary sex differences throughout other regions of the body, in that: “the males of chimpanzee are but
little heavier than the females, and lack such sex characters as cheek pads and huge cranial crests. Even the
sexual distinction in the size of the permanent canine teeth is not nearly as marked and constant in
chimpanzee as in orang-utan and in gorilla. This is shown, e.g., by the data [...] according to which there is
no overlapping of variations in the thickness of the upper canines in males and in females of the latter two
apes, but very considerable over-lapping in chimpanzee (419)”. He concluded that “among the anthropoid
apes, therefore, the amount of specialization in the female pelvis does not correspond to the relative size of
the foetus at birth; it is, however, closely correlated with the general trend for sex differentiation in other
bodily parts, which is extremely well marked in orang- utan and gorilla, but unusually little developed in
gibbon and chimpanzee (422)”.
In a paper entitled “Body Size and Proportions in Chimpanzees, with Special Reference to Pan
troglodytes schweinfurthii from Gombe National Park, Tanzania” Morbeck and Zihlman (1989) undertook a
detailed look at morphological variation in chimpanzees. They used body weight, cranial capacity, linear and
joint area data from a sample of Gombe chimpanzees and compared them to both other Pan troglodytes
populations and Pan paniscus. The Gombe population was represented by 10 adults (6 female, 4 males),
complete and partial skeletons made available through by Goodall. They were compared to 60 adult
specimens (28 females, 22 males) from the Schultz Collection, both captive and wild individuals of mixed
sub-species (22 Pan troglodytes troglodytes, 6 Pan troglodytes verus, and 14 Pan troglodytes
schweinfurthii). Body weights were taken from life measurements and museum records, while other
dimensions were measured from the skeletal material: cranial capacity was estimated by filling the crania
with mustard seeds and measuring the volume, linear measurements (lengths of the scapulae, humeri,
radii, femura, and tibiae, as well as the inonimate length and iliac breadths) were taken with Gneupel sliding
callipers and an osteometric board, while the joint surface areas (glenoid-fossa area, humeral head, the
acetabulum and femoral head areas) were measured from latex casts of dry bones, using a Zeiss MOP3
Analyser, a technique modified from Gomber (1981). Canines were used to confirm the sex diagnoses from
records, measured by Johanson's (1974) method, with the mesio-distal lengths greater than the bucco-
lingual lengths. For the analysis, indices (the humero-femoral, intermembral, brachial, and crural) were
calculated and statistical methods used to find the means, ranges, and standard deviations for each series
of metric variables. “Size adjusted” variables were calculated for each group, for males and females in each
group, and for each recognized subspecies. (372)
23
The results found that the Gombe sample weighed less than other populations, but were similar in
cranial capacity and tooth size. The Gombe chimps were smaller than any other published samples,
including other Pan troglodytes schweinfurthii and Pan paniscus. Cranial capacity was similar to other Pan
troglodytes. While, statistically, the cranial capacity showed no apparent sexual dimorphism, the mean for
the males in the sample was smaller than that of the females, and even smaller than other male averages
from the literature (1989, 373). In terms of dental dimensions, the Gombe sample conformed to the pattern
of variation expected in Pan troglodytes (374). postcranial dimensions, however, were significantly different
and absolutely smaller in most elements: the mean values for the humerus, radius, femur, and tibia lengths,
as well as the forelimb and hindlimb lengths being shorter in the Gombe sample than other Pan troglodytes
(374). Once size-adjusted, these differences disappeared, and the ratios of each bone to the sums of their
elements showed no significant difference from the variation in other samples. The humerus was the
exception to this, with a significant difference between the Gombe sample and captive chimpanzees even in
size-adjusted values (374). The long bone lengths are shorter than Pan troglodytes schweinfurthii and Pan
troglodytes troglodytes, but when compared to Pan troglodytes verus, the means of the humerus and tibia
lengths are similar, whereas the radius and femur means are significantly different. Overall, P. t.
schweinfurthii show the longest lengths, with P. t. troglodytes next, then P. t. verus. The forelimb and
hindlimb segments were shorter in Gombe, but the limb proportions were the same across all populations:
the humero-femoral and intermembral indices were similar between the Gombe sample and other Pan
troglodytes troglodytes, while the brachial and crural indices were comparable in all chimpanzee subspecies
and even bonobos (377).
In linear dimensions of trunk morphology, the average values were intermediate between Pan
troglodytes and bonobos, rather than absolutely or relatively smaller. Clavicular length was significantly
different than other Pan troglodytes, but did not show a significant difference when compared to Pan
paniscus. Iliac breadth was, again, intermediate in absolute size and significantly different from other chimp
subspecies but not from bonobos. (377) The joint surfaces (glenoid-fossa surface area, the humeral head
surface area, and acetabular) were intermediate in size, with the exception of the femoral head surface
area, which was smaller than other subspecies or than bonobos (378). Ratios of joint surface areas within
joints (e.g., the glenoid-fossa to acetabulum, and humeral head to femoral head) show no significant
differences with other subspecies or with Pan paniscus (378).
The comparison between subspecies raises intriguing possibilities as to intra-specific variation;
while sexual dimorphism is only peripherally considered within the paper, the regions of variation, and the
means by which they vary, reveal important areas upon which to focus. The comparison to bonobos is
perhaps somewhat less relevant today than when the paper was initially published (as subsequent work in
24
the fields of both genetics and observational primatology have conclusively confirmed that Pan paniscus
represents a different species); nonetheless, the comparison demonstrates that, even when the absolute
size of some skeletal dimensions enters the range of bonobo size, the shape of elements (as measured by
ratios and indices) remains within the Pan troglodytes pattern.
Relative to other populations, the Gombe chimps seem to be smaller in many but not all ways –
specifically, they present with smaller body weights and long bone lengths, intermediate clavicular and
pelvic measurements, but comparable, if not larger, in dentition and cranial capacities. Morbeck and
Zihlman speculate that “during the process of growth of Gombe chimpanzees which produces overall small
adults, the long bones apparently exhibit greater plasticity. By contrast, joint surface areas and trunk
breadth, reflected in clavicular length and iliac breadth, perhaps come closer to achieving species potential
size” (379). Assuming that the Gombe chimpanzees do, in fact, differ significantly – as opposed to the
appearance of such being an artefact of small sample size in this study, it is too early to even begin to
conjecture as to whether this is a result of developmental plasticity, perhaps triggered by ecological factors,
or is genetic. Nonetheless, the very fact that, as this study clearly demonstrates, some anatomical regions
are more variable than others merits further attention. It is worth noting that the elements that do not
show variation, i.e. cranial capacity and tooth size, are also those which develop earlier in life, whereas the
skeleto-muscular system achieves full growth at a later age – notably, with the full development of
secondary sexual characteristics and sexual dimorphism.
Morbeck published a second paper, focusing specifically on sex differences in the pelvis of Gombe
chimps with known biographic information. With a sample size of 13 (8 females and 5 males),
measurements of pelvic dimensions were modelled after Schultz (1930): ilium length and breadth, ischium
length, pubis length, acetabulum area, and femoral head area (130). Linear lengths were taken with callipers
while joint surface areas were measured from latex templates made from dry bones. In addition, non-
metric data regarding the surface anatomy, i.e. descriptive observations of the dorsal pubis adjacent to the
symphysis and the preauricular area of the ilium (129) were made. While none of the Gombe specimens
demonstrated well-defined resorption areas, there was little visual difference between males and females,
even those known to have given birth, as they showed similar roughness of surfaces (131). Descriptive
statistics were calculated separately for males and females, and the degree of dimorphism presented with
female size as a percentage of male size. The ilium and ischium were found to be longer in males (with 3%
and 4% dimorphism respectively), the pubis longer in females (at -4%), and the acetabulum area relatively
greater in males (11%) – but not to a statistically significant degree. Iliac breadth, however, was significantly
larger in males (at 9%).
Galloway, Morbeck, and Zihlman (1996) looked at sex differences in the vertebral columns of
25
Gombe chimpanzees. Given that chimpanzee sexual dimorphism seems to have only a small presentation in
long bone lengths despite a more significant difference in body weight, which Galloway et al (1996) quote at
24% or greater, alongside the relatively greater joint surface areas in male chimpanzees, they created a
study to investigate the possibility that the skeletal trunk shows significant sexual dimorphism in vertebral
elements. Several scenarios by which these differences could be achieved are hypothesized, (males having a
larger skeletal trunk, or more soft tissue, i.e. muscle mass, or greater bone density).
With 15 adult Pan troglodytes schweinfurthii specimens (8 females and 7 males) compared with
data from Sanders (1994) study of of Pan troglodytes schweinfurthii (3 females and 9 males). Vertebral
elements were measured with digital sliding callipers to the nearest millimetre, with direct areal
measurements obtained from latex moulds of the superior surface of vertebral centra, and digitized using a
Zeiss-MOP 3-image analyser. (444). Statistical analysis of the results revealed some significant sex
differences. Male vertebral bodies throughout the spinal column are larger than those of females, as
indicated by both linear and surface measurements (452). Significant differences with sex occur in linear
measurements of the spine, while sex differences in segment length were minimal and not statistically
significant. Direct measurements showed statistically significant differences in the last cervical vertebrae
and almost all thoracic and lumbar vertebrae; in addition, segment lengths “suggest possible differences
associated with sex in vertebral column curvature” (452).
In line with Morbeck and Zihlman's (1989) finding that Gombe chimpanzees were lighter in body
mass, despite having cranial capacities, tooth sizes, and limb proportions similar to other Pan troglodytes,
the Gombe sample demonstrated vertebral measurements that were considerably smaller than previously
published values. Surface areas of vertebral centra were smaller than those reported by Shapiro (1991) or
Sanders (1994). (451) However, the degree of sexual dimorphism within the Gombe sample is “much more
pronounced, particularly in vertebral breadth and surface measurements” (452). Galloway attributes this to
the presence of absolutely smaller females in the Gombe sample. Furthermore, the importance of the
gleno-humeral articulation in weight-bearing and the sexual dimorphism it displays in favour of males,
seems that the size difference reflects a greater need for support of the males' larger body, particularly as
other load-bearing surfaces through the vertebral column are similarly enlarged. This data, when
contextualized with previous studies, leads to the conclusion that in chimpanzees, the male torso is only
slightly longer, but much wider (Galloway et al 1992, 452). Galloway further conjectures that these
differences are developmental; the possibility of differing locomotor strategies between the sexes is raised,
although he notes that not only would further study on positional behaviour and anatomy be required to
fully consider such a possibility, but a better understanding of size-related sex differences is needed first
(453) .
26
In 2008, Zihlman and several co-authors published a study continuing, and directly playing off, the
work that had begun with Morbeck (1989): “Morphological variation in Adult Chimpanzees (Pan troglodytes
verus) of the Tai National Park, Cote D'Ivoire”. Unlike Zihlman's previous work with Morbeck, this 2008 study
did directly consider sexual dimorphism. A new sample of 25 Pan troglodytes verus (15 females and 10
males), collected over 19 years in the field as part of a long-term observational study, was assessed and
compared to the Pan troglodytes schweinfurthii measurements from earlier studies (2008, 34). Cranial
capacity and postcranial dimensions were taken in the same manner as the previous studies, and included
scapular and clavicular lengths, innonimate length, iliac breadth, and acetabular diameter. A number of
cranial-facial measurements were added, following landmarks as defined by Cramer (1977), taken with
spreading and digital callipers (34). The cranial sample size was larger than that for the postcranial
dimensions, as 10 immature specimens (7 female and 3 male) were added; as the second molars in these
specimens had already erupted, and brain size is nearly complete by this time, they were sufficiently mature
(35). For the rest of the sample specimens, maturity was defined as the eruption and occlusion of all teeth,
and the complete fusion of the epiphyses of proximal humeri (35).
Indices were calculated, and all elements were analysed statistically by sex within the Tai group, by
sex within the Gombe group, by combined sex samples, and between the sexes of each group (36). While
individual measurements were not presented, the sample sizes, means, and ranges of measurements were
given with p-values and F-values; sexual dimorphism of specific dimensions were calculated; the adult
female means as a percentage of the adult male means, as opposed to the percentage difference that had
been presented in Morbeck and Zihlman (1989). Clavicular length was significantly sexually dimorphic
within the Tai sample (92.0%), but not for the Gombe sample (91.7%), while the sexual dimorphism of the
scapular length was significantly different in both samples (at 87.8% and 87.5% respectively). Iliac breadth,
inonimate length, and acetabulum diameter were not significant within either the Tai or Gombe samples –
yet, the combined sex samples of all three dimensions did show significant sexual dimorphism, and the
differences between Tai and Gombe females was also highly significant. (2008, 38)
Sex differences were found to be mosaic between sites. Tai males and females differed in clavicular
and scapular dimensions, but not in pelvic morphology. The difference in scapular breadth common to Tai
and Gombe confirmed earlier observations, by agreeing with Galloway et al (1996), who found that the last
cervical and almost all thoracic and lumbar vertebrae were dimorphic in the Gombe sample. Tai females
accounted for difference in cranial capacity between sites, as means and ranges of males from both sites
were nearly identical, yet Tai females and males differed significantly. These results differed from previous
findings regarding cranial capacity and dimorphism in Pan troglodytes verus (Dierbach 1986) and, those
from another study of the same collection, by Protsch von Zieten (1987); however, given that Zihlman's
27
(2008) had a much larger sample size and considered more facial elements, this is neither surprising nor
necessarily worrisome. The limb indices were not found to differ significantly between Tai and Gombe,
although the bone lengths themselves did differ. Most of the difference between sites was due to the
females: Gombe females had higher cranial capacities, whereas Tai females had longer limbs and larger
pelves. However, the size of different features did not seem to necessarily correlate (i.e., a larger than
average cranium did not predict larger long bone lengths); in fact, a Tai female with the smallest cranial
capacity had above average limb measurements, while a Gombe male with the shortest limb lengths has a
large cranial capacity. “Morphological distinctions in the pattern of sexual dimorphism of the Tai sample is
not inconsistent with findings based on other morphological and genetic studies. The morphological
findings reported here may be an artefact of sampling, may reflect genetic drift or adaptation to particular
ecological conditions, or reflect some combination thereof. (40) “
28
Table 1: Data from previously published studies
Morbeck &
published: Morbeck et al 1992
29
Zihlman et al 2008 Zihlman1989

Schultz 1937
population: Gombe Gombe
Tai Gombe
P. t.
subspecies: P. t. verus P. t. schweinfurthii
P. t. schweinfurthii schweinfurthii
Summary
Summary Sexual dimorphism Summary statistics Sexual dimorphism Sexual dimorphism Summary statistics
statistics
statistics
female female female
female
significance mean significance significance mean as mean as
N mean N mean mean as N mean N mean
level as % level level % of % of
% of male
of male male male
scapula breadth 80.200
M 5 < 0.05 87.80% M 3 83.4000 <0.01 87.50%
0
70.200
F 10 F 8 73.0000
0
Total Total 11 78.2000
scapula length M M
F F
Total Total
clavicle length 121.30 116.600
M 3 < 0.05 92.00% M 5 91.70%
00 0
111.50 107.000
F 9 F 8
00 0
116.30 111.800
Total 12 Total 13 95.90% 9 109.4
00 0
inonimate length 271.90 263.700
M 6 96.80% M 5
00 0
263.30 252.800
F 7 F 8
00 0
267.60 258.200
Total 13 Total 13 8 254.9
00 0
ilium length M -- M 97.00%
F F
Total Total
ilium breadth 121.80 114.000
M 5 93.00% M 6 92.30% 8 104 <0.05 91.00%
00 0
113.30 104.100
F 7 F 8
00 0
109.000
Total 12 Total 14
0
ischium length M -- M 96.00%
F F
Total Total
pubis length M -- M 104.00%
F F
Total Total
acetabulum 39.100
M 6 96.70% M 5 35.9000 94.50%
diameter 0
published: Zihlman et al 2008 Morbeck & Morbeck et al Schultz1937
Zihlman1989 1992
297.90 277.00 306.00
humerus length M 5 99.60% M 6 98.10% 89.00% M 45 97.71%
30
00 00 00
296.80 271.00 299.00
F 9 F 8 F 62
00 00 00
297.40 274.40
Total 14 Total 14 14 301.6 Total
00 00
Total Total Total
272.10 256.80
radius length M 4 98.80% M 5 95.90% M
00 00
268.90 246.30
F 9 F 8 F
00 00
251.60
00
297.00 274.30 256.00
femur length M 6 97.30% M 4 97.30% M 38 97.27%
00 00 00
288.90 266.80 249.00
F 8 F 7 F 58
00 00 00
270.50
00
femural head area M M 97.00% M
F F F
Total Total Total
243.30 230.80
tibia length M 6 98.50% M 5 96.50% 14 254.5 M
00 00
239.60 222.70
F 7 F 7 F
00 00
226.70
Total 13 Total 12 Total
00
intermembral 107.30
M M 7 106 M 44 99.16%
index 00
106.40
F F F 60
00
105.90
Total 106.7 Total Total
00
humeral-femoral
M M M
index
F F F
101.40
Total 102.2 Total 7 102 Total
00
93.100
brachial index M M M 45 99.46%
0
92.600
F F F 60
0
91.800
Total 90 Total 9 91 Total
0
83.500
crural index M M M 44 99.88%
0
83.400
F F F 60
0
83.800
Total 81.9 Total 7 83 Total
0
Methods
Sample
The specimens used in this study are from the Powell-Cotton Museum in Quex Park, Kent. The
specimens were collected in Cameroon between 1927 and 1935. They were collected by Fred Merfield and
local villagers in hunting expeditions; cause of death was presumably ballistic trauma in all cases (many of
the specimens had clear bullet wounds in the scapulae) and the remains had been transported to the
United Kingdom shortly after death.
A total of 56 specimens (36 female, 20 male) are considered in this study. Only 4 specimens (3
female, 1 male) lacked a complete fusion of the epiphyses, but these were seemingly full-grown and of
comparable size to the rest of the sample. The sex of all specimens had been recorded at the time of death
and confirmed in previous cranial studies of the collection. Sample sizes for specific dimensions and
elements differ, as some specimens suffered from post-mortem damage and/or showed pathological
abnormalities (including healed fractures of the femora and clavicles) and had to be excluded; in a few
cases, right elements were substituted for the left when the latter was too damaged or abnormal to allow
accurate measurements. In situations in which elements were in multiple pieces, but did not seem to lack
material, measurements were taken, and the fact that they represented approximations noted.
Data collection
Linear measurements of postcranial elements (including the scapulae, clavicles, vertebrae, humeri,
radii, ulnae, ossa coxae, femora, and tibiae) were taken for the left sides of all 56 specimens. Measurements
of both left and right elements were taken on a subset of 36 specimens (18 female, 18 male), allowing for
the consideration of bilateral asymmetry. Maximum lengths of all long bones (humeri, ulnae, radii, femora,
and tibiae) and the ossa coxae were taken with an osteometric board and recorded to the nearest
millimetre; all other dimensions were taken with sliding digital callipers and recorded to the nearest
hundredth of a millimetre. All measurements were taken by one researcher, myself, over the course of a
single week while on site at the Powell-Cotton Museum.
Measurements were chosen with reference to the literature on chimpanzee, primate, and human
sexual dimorphism. The intention was to extend previous work by increasing the number of anatomical
regions considered within a single study while simultaneously expanding the number of measurements
taken on each skeletal element so as to better examine shape in addition to size. Specific measurements
were adopted from Jay Stock's protocols for measuring human remains. Matching all measurements and
31
landmarks as closely as possible allowed for direct comparison with human populations previously
measured by Jay Stock. For this reason, much of the terminology used is common to the study of human
anatomy as opposed to the field of zoology. This choice, given the lack of contemporary work looking
chimpanzee postcranial anatomy in comparable detail, the great similarity of the Pan troglodytes skeletal
anatomy to Homo sapiens, and the interest in juxtaposing their morphologies when interpreting the
hominid fossil record, seemed appropriate.
While previous studies have focused primarily on the femur and pelvic regions, I chose to broaden
the scope of this study and include at least basic dimensions of all limb elements. In order to approximate
robusticity, further dimensions (including mid-shaft, head, and neck diameter) were measured. As previous
work indicates, it is in torso girth – as opposed to height or even weight – that living chimpanzees are
particularly dimorphic, particular attention, by means of more detailed measurements, was paid to the
humerus, scapula, and clavicle. Crania were not considered in this study as they have been, until this point
in time, the most thoroughly researched. However, if this methodology were to extend to consider intra-
specific variation and compare between populations, it would be of great interest to include cranial
dimensions as there is reason to believe that cranial dimorphism may not scale with dimorphism
throughout the rest of the chimpanzee body.
Field notes by Fred Merfield from the expeditions during which the specimens were collected were
made available by the Powell-Cotton Museum. These field notes documented various biometric
measurements, descriptions of the animals' appearance and health, as well as conjecture as to their sub-
classification. Biometric data was available for a subset of 29 specimens (18 females, 11 males). Weight,
which was recorded in only three relevant cases, was discarded as the method by which it was measured (of
the entire animal, of the corpse lacking the skin or other organs) was inconsistent. Height and span were
available for this entire subset of 29 specimens; head-to-fork length, girth circumference, chest
circumference, belly circumference, bicep circumference, forearm circumference, neck circumference, hand
length and width, and foot length and width dimensions were recorded in various combinations thereafter.
Analysis
Data are analysed using the Statistical Package for the Social Sciences.
(1) Indices and ratios:
From the measurements, a variety of proportions and indices were calculated, including the intermembral
index, the brachial index, and the crural index, and the claviculo-humeral index. Numerous morphological
values were estimated, including forelimb and hindlimb lengths, from the skeletal measurements. For each
skeletal element, ratios of each measurement against the total length of the bone as well as ratios of several
32
of the measurements to each other, were calculated, to provide size-adjusted values and to allow
consideration of proportional differences.
(2) Evaluating sexual dimorphism within the population:
Sample sizes, means, standard deviations, and ranges were calculated for the entire sample and
independently by sex. The means by sex were compared using an independent samples t-test. The
measurements which demonstrated statistically significant sex differences between female and male means
are considered in further detail and ratios of sexual dimorphism calculated as the female mean as a
percentage of male mean.
(2) Comparing body proportions:
The biometric data (external body dimensions from Merfield's field notes) were treated to the same
statistical analyses as the measurements taken from the Powell-Cotton collection.
(3) Predicting sex:
Discriminant analysis was used to test the ability of the significantly dimorphic measurements to predict sex
classification. Models were based on (a) multiple dimensions of a single skeletal element (e.g.: scapula,
clavicle, pelvis, humerus, radius, femur and tibia); (b) multiple skeletal elements within an anatomical
feature (e.g. dimensions of forelimb elements, dimensions of hindlimb elements, dimensions of the
shoulder girdle, dimensions of the pelvis, and dimensions of the trunk); and (c) ratios reflecting
proportional and size-adjusted values.
33
Results
Summaries of all measurements, with statistical descriptives (including sample sizes, means,
standard deviations, coefficients of variation, minimum, and maximum values) by sex and for the
population total are presented alongside the results of independent samples t-test (t-values, degrees of
freedom, and p-values) compare with the sex means, in Tables 2-6. The same information is presented for
indices and ratios which comparing separate skeletal elements in Table 7, and for a variety of size-adjusted
and proportional dimensions in Table 8.
Results concerning sex differences are organized by the following anatomical regions: shoulder
girdle, pelvis forelimb, and hindlimb. Tables presenting information are interspersed with the text, arranged
by skeletal element, with the statistical significance of each dimension listed alongside the sexual
dimorphism (calculated as female mean as a percentage of male mean). The population distributions by sex
of each significantly dimorphic measurement are displayed in graphs. The results of the discriminant
function analysis are presented after the skeletal element or anatomical region from which they were
derived.
Statistical descriptives of the biometric information regarding external body dimensions is
presented in Table 9, and the result of a final discriminant function analysis using only the proportional
measurements to classify by sex is presented in Table 28.
34
Table 2: Measurements of the scapula

Independent samples t-test
Summary statistics (between sexes)
N mean std CV minimum maximum t-value df p-value
scapula length
M 17 152.49 7.2949 0.0478 137.43 170.50 2.4270 51 0.0190
F 36 145.71 10.3347 0.0709 123.42 177.60
Total 53 147.88 9.9224 0.0671 123.42 177.60
scapula breadth M 18 86.85 3.6347 0.0419 78.03 91.13 3.0040 52 0.0040
F 36 81.94 6.4110 0.0782 56.43 96.33
Total 54 83.58 6.0685 0.0726 56.43 96.33
glenoid fossa (max) length M 18 38.14 1.9066 0.0500 35.02 42.88 2.6790 45.3570 0.0100
F 36 36.45 2.6618 0.0730 31.72 41.82
Total 54 37.02 2.5481 0.0688 31.72 42.88
glenoid fossa breadth M 18 24.85 1.9736 0.0794 22.18 29.16 2.1830 52 0.0340
F 36 23.64 1.8988 0.0803 19.71 28.41
Total 54 24.04 1.9907 0.0828 19.71 29.16
spine length M 16 142.32 6.7841 0.0477 129.65 152.31 1.7130 42 0.0940
F 28 138.20 8.1084 0.0587 121.48 157.79
Total 44 139.70 7.8320 0.0561 121.48 157.79
spine height M 16 28.09 2.5898 0.0922 24.24 32.59 0.9900 43 0.3280
F 29 27.11 3.4548 0.1274 20.72 38.65
Total 45 27.46 3.1792 0.1158 20.72 38.65
coracoid length M 16 41.36 2.9710 0.0718 35.67 45.67 0.9230 43 0.3610
F 29 40.60 2.4616 0.0606 34.57 46.00
Total 45 40.87 2.6459 0.0647 34.57 46.00
acromion breadth M 16 21.96 3.1694 0.1443 19.21 32.47 1.3130 42 0.1960
F 28 20.59 3.3997 0.1651 15.89 32.47
Total 44 21.09 3.3471 0.1587 15.89 32.47
superior border length M 16 28.49 4.8420 0.1700 20.84 38.81 1.3700 43 0.1780
F 29 26.16 5.7901 0.2213 13.17 36.09
Total 45 26.99 5.5323 0.2050 13.17 38.81
35
Table 3: Measurements of the clavicle

Summary statistics
(between sexes)
clavicle maximum length M 19 129.29 5.9945 0.0464 117.31 139.81 2.6640 51 0.0100
F 35 124.31 6.6619 0.0536 109.29 137.69
Total 53 126.00 6.8146 0.0541 109.29 139.81
mid-shaft diameter (maximum) M 18 13.24 1.7557 0.1326 11.27 19.24 3.2660 52 0.0020
F 36 11.87 1.2855 0.1083 9.78 14.77
Total 54 12.33 1.5833 0.1284 9.78 19.24
mid-shaft diameter (minimum) M 19 8.64 0.9032 0.1045 7.13 10.95 3.3240 52 0.0020
F 35 7.75 0.9387 0.1211 6.1 10.02
Total 54 8.05 1.0113 0.1256 6.1 10.95
conoid tubercle superior- M 18 17.19 2.2365 0.1301 13.16 20.4 0.0820 52 0.9350
inferior diameter F 36 17.13 2.8660 0.1673 11.54 22.12
Total 54 17.15 2.6514 0.1546 11.54 22.12
conoid tubercle anterior- M 18 8.53 1.0927 0.1281 6.35 10.3 3.6780 52 0.0010
posterior diameter F 36 7.39 1.0644 0.1440 5.57 9.73
Total 54 7.77 1.1939 0.1537 5.57 10.3
sternal facet height (maximum) M 18 22.46 1.5752 0.0701 18.74 24.56 2.1370 52 0.0370
F 36 21.26 2.1019 0.0989 16.32 25.52
Total 54 21.66 2.0099 0.0928 16.32 25.52
sternal facet breadth M 18 14.89 2.4720 0.1660 8.2 18.38 0.7270 52 0.4710
(minimum)
F 36 14.41 2.2407 0.1555 9.51 19.28
Total 54 14.57 2.3085 0.1584 8.2 19.28
36
Table 4: Measurements of the pelvis

Independent samples t-
Summary statistics
test (between sexes)
os coxae height M 18 276.06 10.65 0.0386 257.00 296.00 1.762 51 0.084
F 35 270.06 12.2473 0.0454 243.00 293.00
Total 53 272.09 11.9748 0.0440 243.00 296.00
ilium breadth M 18 124.74 9.9170 0.0795 105.07 144.70 2.330 51 0.024
F 35 118.39 9.1149 0.0770 101.71 136.83
Total 53 118.85 17.5058 0.1473 13.81 144.70
Iliac crest (max) breadth M 18 7.79 1.3126 0.1685 5.94 10.32 1.168 52 0.248
F 35 7.26 1.6919 0.2330 4.38 10.17
Total 54 7.43 1.5834 0.2131 4.38 10.32
Iliac crest (min) breadth M 18 3.53 0.7033 0.1992 2.54 4.70 4.268 52 0.000
F 35 2.76 0.5764 0.2088 1.79 4.00
Total 54 3.02 0.7145 0.2366 1.79 4.70
acetabular height M 19 39.57 2.8843 0.0729 36.08 48.04 3.789 53 0.000
F 36 36.29 3.1334 0.0863 28.72 45.17
Total 55 37.42 3.4077 0.0911 28.72 48.04
acetabulosciatic breadth M 19 29.77 1.6932 0.0569 26.51 32.33 1.942 51.3780 0.058
F 36 28.27 3.9787 0.1407 21.41 39.71
Total 55 28.79 3.4247 0.1190 21.41 39.71
superior pubic height M 19 15.14 1.6466 0.1088 10.87 17.96 -0.060 53 0.951
F 36 15.17 1.9810 0.1306 11.28 20.54
Total 55 15.16 1.8567 0.1225 10.87 20.54
symphyseal height M 18 51.86 9.3108 0.1795 40.39 70.50 1.893 23.28 0.071
F 36 47.35 5.5738 0.1177 39.06 60.07
Total 54 48.85 7.2754 0.1489 39.06 70.50
symphyseal breadth M 18 11.46 2.0431 0.1783 8.03 15.89 2.412 52 0.019
F 36 9.88 2.3775 0.2406 6.01 15.03
Total 54 10.40 2.3747 0.2283 6.01 15.89
auricular surface length M 19 52.30 6.2292 0.1191 44.68 66.47 2.694 53 0.009
F 36 47.90 5.5104 0.1150 37.88 57.29
Total 55 49.42 6.0894 0.1232 37.88 66.47
auricular surface breadth M 19 15.31 2.7697 0.1809 9.55 20.38 1.904 53 0.062
F 36 13.99 2.2367 0.1599 8.16 17.75
Total 55 14.45 2.4892 0.1723 8.16 20.38
as
37
Table 5: Measurements of the humerus, ulna, and radius

Summary statistics
(between sexes)
humerus length M 19 300.89 11.2640 0.0374 285.00 320.00 0.5290 53 0.5990
F 36 298.94 13.8047 0.0462 261.00 322.00
Total 55 299.62 12.9107 0.0431 261.00 322.00
humerus mid-shaft diam (max) M 19 24.89 1.7735 0.0713 21.45 28.70 2.4510 53 0.0180
F 36 23.69 1.7075 0.0721 20.02 27.33
Total 55 24.11 1.8086 0.0750 20.02 28.70
humerus mid-shaft diam (min) M 19 21.81 1.4780 0.0678 18.88 24.98 1.9430 53 0.0570
F 36 20.74 2.1580 0.1041 17.15 28.39
Total 55 21.11 2.0033 0.0949 17.15 28.39
humerus 35% diam (max) M 19 25.35 2.1384 0.0844 20.58 29.20 1.8310 52 0.0730
F 35 24.30 1.9259 0.0793 20.70 28.15
Total 54 24.67 2.0460 0.0829 20.58 29.20
humerus 35% diam (min) M 19 22.19 1.9019 0.0857 19.15 25.73 2.4250 52 0.0190
F 35 20.92 1.8009 0.0861 17.05 24.04
Total 54 21.37 1.9192 0.0898 17.05 25.73
humerus proximal transverse M 19 47.01 2.3928 0.0509 43.06 51.02 3.5280 53 0.0010
diameter F 36 43.36 4.1506 0.0957 24.40 48.92
Total 55 44.62 4.0180 0.0900 24.40 51.02
humerus proximal head diam M 19 41.71 2.9749 0.0713 37.03 47.24 4.1660 29.8 0.0000
F 36 38.44 2.3169 0.0603 33.45 43.26
Total 55 39.57 2.9811 0.0753 33.45 47.24
humerus surgical neck diam M 19 28.47 4.0635 0.1427 21.13 41.35 3.8910 53 0.0000
(max) F 36 25.19 2.2103 0.0877 20.85 30.95
Total 55 26.32 3.3387 0.1269 20.85 41.35
humerus surgical neck diam M 19 23.05 2.2221 0.0964 18.74 27.90 4.1440 53 0.0000
(min) F 36 20.67 1.9115 0.0925 17.03 24.10
Total 55 21.49 2.3054 0.1073 17.03 27.90
humerus epicondylar breadth M 18 63.88 2.8765 0.0450 58.39 69.54 4.3440 52 0.0000
F 36 59.42 3.8443 0.0647 52.31 67.72
Total 54 60.90 4.1131 0.0675 52.31 69.54
humerus distal articular breadth M 18 47.93 4.4943 0.0938 43.49 63.15 3.5910 52 0.0010
F 36 44.56 2.4320 0.0546 38.24 49.86
Total 54 45.68 3.5999 0.0788 38.24 63.15
ulna length (max) M 19 294.89 11.2788 0.0382 275.00 318.00 2.3330 53 0.0230
F 36 285.08 16.3609 0.0574 240.00 320.00
Total 55 288.47 15.4295 0.0535 240.00 320.00
radius length (max) M 19 279.53 11.4036 0.0408 260.00 302.00 2.3000 53 0.0250
F 36 270.58 14.7559 0.0545 237.00 305.00
Total 55 273.67 14.2440 0.0520 237.00 305.00
radius mid-shaft diam (max) M 19 16.23 1.5190 0.0936 13.90 19.21 2.7900 53 0.0070
F 36 14.97 1.6411 0.1096 11.36 18.79
Total 55 15.40 1.6982 0.1103 11.36 19.21
radius mid-shaft diam (min) M 19 13.80 1.0619 0.0769 11.79 15.30 2.4570 53 0.0170
F 36 13.00 1.1968 0.0921 10.48 15.47
Total 55 13.28 1.2053 0.0908 10.48 15.47
radius head diam M 19 25.86 1.6879 0.0653 23.51 29.47 4.3300 53 0.0000
F 36 23.85 1.5995 0.0671 20.34 26.41
Total 55 24.55 1.8790 0.0765 20.34 29.47
38
Table 6: Measurements of the femur and tibia

Summary statistics
(between sexes)
femur length M 18 296.56 12.9489 0.0437 276.00 315.00 0.8950 52 0.3750
F 36 292.64 16.1195 0.0551 255.00 323.00
Total 54 293.94 15.1277 0.0515 255.00 323
femur mid-shaft diam (max) M 19 26.41 1.0699 0.0405 24.53 28.46 3.1520 51 0.0030
F 34 24.94 1.8712 0.0750 21.63 30.32
Total 53 25.47 1.7687 0.0694 21.63 30.32
femur mid-shaft diam (min) M 19 22.47 1.6571 0.0737 19.41 25.53 1.3100 51 0.1960
F 34 21.87 1.5637 0.0715 19.28 26.18
Total 53 22.08 1.6082 0.0728 19.28 26.18
femur head diam (max) M 19 34.20 2.1264 0.0622 31.24 40.41 3.0400 53 0.0040
F 36 32.44 2.0016 0.0617 28.96 37.03
Total 55 33.05 2.1954 0.0664 28.96 40.41
femur SI height of the M 19 23.39 1.9360 0.0828 18.54 25.88 1.9840 53 0.0530
femoral neck F 36 22.22 2.1295 0.0958 18.18 26.17
Total 55 22.62 2.1212 0.0938 18.18 26.17
femur neck AP diam M 19 19.69 1.8378 0.0933 16.30 23.24 1.4700 53 0.1480
F 36 18.87 2.0465 0.1085 15.73 24.68
Total 55 19.15 1.9992 0.1044 15.73 24.68
femur neck length (min) M 19 24.43 2.4670 0.1010 20.26 29.19 2.1770 53 0.0340
F 36 22.65 3.0644 0.1353 16.04 28.04
Total 55 23.26 2.9734 0.1278 16.04 29.19
femur biomechanical neck M 19 68.17 2.8620 0.0420 64.11 74.78 6.8490 53 0.0000
length F 36 62.81 2.7023 0.0430 56.14 67.59
Total 55 64.66 3.7508 0.0580 56.14 74.78
femur epicondylar breadth M 19 64.51 3.5742 0.0554 57.94 70.80 5.1390 53 0.0000
F 26 59.48 3.3870 0.0569 51.49 66.86
Total 55 61.22 4.1858 0.0684 51.49 70.80
femur distal anterior- M 19 26.97 1.5169 0.0562 25.07 30.32 2.1920 51 0.0330
posterior diam F 34 25.86 1.8830 0.0728 22.15 29.23
Total 53 26.26 1.8258 0.0695 22.15 30.32
tibia length (max) M 19 252.89 10.4663 0.0414 232.00 273.00 1.1760 53 0.2450
F 36 248.36 14.9593 0.0602 220.00 283.00
Total 55 249.93 13.6488 0.0546 220.00 283.00
tibia mid-shaft (ap) diam M 19 23.73 1.5162 0.0639 21.33 26.99 2.8340 53 0.0060
(max) F 36 22.15 2.1541 0.0973 18.74 27.61
Total 55 22.69 2.0846 0.0919 18.74 27.61
tibia mid-shaft diam (min) M 19 15.73 1.2362 0.0786 12.86 17.81 2.0190 53 0.0490
F 36 14.85 1.6705 0.1125 12.63 19.76
Total 55 15.15 1.5800 0.1043 12.63 19.76
plateau breadth M 18 60.24 2.8026 0.0465 56.22 65.84 5.6560 52 0.0000
F 36 55.38 3.0534 0.0551 49.52 62.02
Total 54 57.00 3.7435 0.0657 49.52 65.84
39
Table 7: Limb lengths, ratios, and indices

Independent samples t-
Summary statistics
test (between sexes)
N mean std CV t-value df p-value
forelimb length M 19 580.421053 21.841133 0.0376 1.5 53 0.140
F 36 569.527778 27.381418 0.0481
Total 55 573.290909 25.928443 0.0452
hindlimb length M 18 549.666667 22.955199 0.0418 1.07 52 0.290
F 36 541.000000 30.362571 0.0561
Total 54 543.888889 28.192510 0.0518
clavicle–humerus M 18 43.004479 2.521794 0.0586 1.94 51 0.060
index F 35 41.699006 2.212249 0.0531
Total 53 42.142374 2.380879 0.0565
clavicle-femur index M 17 43.734241 2.686088 0.0614 1.63 50 0.110
F 35 42.626085 2.086718 0.0490
Total 52 42.988366 2.332808 0.0543
clavicle–forelimb ratio M 18 22.255527 1.279425 0.0575 1.06 51 0.300
F 35 21.886032 1.164402 0.0532
Total 53 22.011521 1.205348 0.0548
humero-femoral index M 18 101.617672 2.951126 0.0290 -0.81 52 0.420
F 36 102.226559 2.407073 0.0235
Total 54 102.023597 2.589144 0.0254
brachial index M 19 92.907083 1.980777 0.0213 3.43 53 0.000
F 36 90.513948 2.676743 0.0296
Total 55 91.340667 2.696432 0.0295
crural index M 18 85.373411 1.820480 0.0213 0.86 52 0.400
F 36 84.872333 2.115271 0.0249
Total 54 85.039359 2.018576 0.0237
intermembral index M 18 105.737593 2.707592 0.0256 0.52 52 0.610
F 36 105.345809 2.591553 0.0246
Total 54 105.476404 2.610000 0.0247
40
Table 8: Sexual dimorphism in proportions

summary statistics Independent samples t-test
(between sexes)
ratio of:
N mean std CV t-value df p-value
clavicle mid-shafter M 18 0.102576 0.014472 0.1411 2.127 51 0.038
diameter (max) to
clavicle length F 35 0.095109 0.010727 0.1128
Total 53 0.097645 0.012508 0.1281
acetabular diameter to M 18 0.143167 0.009800 0.0685 2.932 51 0.005
os coxae height
F 35 0.134361 0.010621 0.0790
Total 53 0.137352 0.011085 0.0807
symphyseal breadth to M 18 0.041526 0.007290 0.1756 2.110 51 0.040
os coxae height
F 35 0.036447 0.008762 0.2404
Total 53 0.038172 0.008571 0.2245
humerus proximal M 19 0.156408 0.009322 0.0596 -0.997 53 0.001
diameter by humerus
length F 36 0.145092 0.012877 0.0888
Total 55 0.149001 0.010681 0.0717
humerus epicondylar M 18 0.212301 0.010681 0.0503 4.062 52 0.000
breadth to humerus
length F 36 0.198898 0.011775 0.0592
Total 54 0.203365 0.012993 0.0639
femur biomechanical M 18 0.230796 0.008255 0.0358 5.756 51 0.000
neck length
F 35 0.214950 0.010053 0.0468
Total 53 0.220332 0.012073 0.0548
femur epicondylar M 18 0.217994 0.013170 0.0604 4.106 52 0.000
breadth to femur length
F 36 0.204092 0.010960 0.0537
Total 54 0.208726 0.013369 0.0641
tibia plateau breadth by M 18 0.238433 0.011768 0.0494 3.703 52 0.001
tibia length
F 36 0.223527 0.014889 0.0666
Total 54 0.228496 0.015528 0.0680
41
Table 9: Biometric data from field notes, Merfield 1933
Summary statistics Independent samples t-test

(between sexes)
female mean
as % of
N mean std CV t-value df p-value male
height M 9 47.36 2.8644 0.0605 -1.013 28 0.320 101.79%
F 21 48.21 1.7235 0.0357
Total 30 47.96 2.1143 0.0441
span M 9 69.71 3.4585 0.0496 0.149 28 0.883 99.71%
F 21 69.51 3.2629 0.0469
Total 30 69.57 3.2635 0.0469
head-to-
M 6 31.69 2.2273 0.0703 0.308 22 0.761 99.27%
fork
F 18 31.46 1.3284 0.0422
Total 24 31.52 1.5470 0.0491
girth M 4 39.88 10.0799 0.2528 0.757 3.09 0.503 90.35%
F 15 36.03 2.3354 0.0648
Total 19 36.84 4.8750 0.1323
chest M 5 37.10 2.3022 0.0621 0.997 9 0.345 94.96%
F 6 35.23 3.6111 0.1025
Total 11 36.08 3.0975 0.0859
belly M 5 35.90 6.6370 0.1849 0.112 19 0.257 107.33%
F 16 38.53 3.5624 0.0925
Total 21 37.90 4.4325 0.1170
biceps M 7 11.29 1.6421 0.1454 0.029 24 0.977 99.82%
F 19 11.27 1.0674 0.0947
Total 26 11.27 1.2114 0.1075
forearm M 6 11.04 1.1770 0.1066 0.119 23 0.907 99.55%
F 19 10.99 0.9269 0.0843
Total 25 11.00 0.9662 0.0878
neck M 8 15.47 1.7343 0.1121 1.934 24 0.065 93.02%
F 18 14.39 1.0955 0.0761
Total 26 14.72 1.3844 0.0940
hand
M 8 9.12 0.7260 0.0796 -0.631 25 0.534 101.75%
height
F 19 9.28 0.5695 0.0614
Total 27 9.23 0.6102 0.0661
foot height M 9 9.44 0.5966 0.0632 0.159 25 0.875 99.68%
F 18 9.41 0.5033 0.0535
Total 27 9.42 0.5248 0.0557
42
Shoulder girdle
In all measured dimensions of the scapula,
Table 10: Sex differences in the scapula
the male mean is larger than the female. While in
significance female mean as %
some cases the female maximum exceeds the male level of male mean
scapula length < 0.05 95.56%
(scapula length and breadth, spine length and
scapula breadth < 0.01 94.35%
height, and coracoid length), the female mean is glenoid fossa length 0.01 95.56%
nonetheless smaller than both the combined sex glenoid fossa breadth < 0.05 95.12%
spine length – 97.11%
mean and the male mean. The differences are
spine height -- 96.51%
minimal, as are those in the other dimensions, coracoid process length -- 98.16%
ranging from 98.16% dimorphic (coracoid length) acromion breadth -- 93.78%

superior border length -- 91.80%
to 91.80% (superior border length). Only a few
measurements have statistically significant sex
differences: the scapula lengths (male 152.49 mm, N = 17 ; female 145.71 mm, N = 36) and breadths (male
86.85 mm, N=18; female 81.94, N=36) are dimorphic, with females averaging 95.56% (p = 0.019) and
94.35% (p = 0.004), respectively. The glenoid fossa is significantly dimorphic in both length (male 38.14 mm,
N = 18 ; female 36.45 mm, N = 36) and breadth (male 24.85 mm, N=18; female 23.64, N=36) at 95.56% (p =
0.010) and 95.12% (p = 0.034).
Table 11: Scapular Model: discriminant function analysis
Eigenvalues Wilks' Lambda Standardized canonical Classification

discriminant function results
coefficients
% of Wilks'
Eigenvalue variance Lambda Chi- sq df sig male female
0.341 100.00% 0.746 14.364 4 0.006 scapula length 0.674 82.40% 69.40%
glenoid fossa length 0.290 Total: 75.90%
glenoid fossa breadth 0.182
Using the significant scapular dimensions to predict sex of an individual resulted in a model which
was better able to identify males than females. The scapula breadth contributed the most to the
identification, followed closely by the scapula length. The dimensions of the glenoid fossa contributed to
relatively small degrees, despite their comparable levels of dimorphism at similar degrees of statistical
significance.
43
Figure 1: Scapula lengths by sex

Figure 2: Scapula breadths, by sex
Figure 3: Glenoid fossa length Figure 4: Glenoid fossa breadth
44
In all measured dimensions of the

Table 12: Sex differences in the clavicle
clavicle, the male mean exceeds the significance female mean as % of
level male mean
female. All measurements show relatively
clavicle maximum length 0.01 96.15%
similar levels of dimorphism, meaning that clavicle mid-shaft diam (max) < 0.01 89.63%
the dimensions seem to scale with size. clavicle mid-shaft diam (min) < 0.01 89.71%
conoid tuberclue SI diam (min) -- 99.63%
The clavicle shows statistically significant clavicle maximum (AP) diameter at
<0.01 86.64%
conoid tubercle
sex differences in all measurements taken 94.66%
clavicle sternal facet (max) height < 0.05
with the exception of two: the superior- clavicle sternal facet (min) breadth -- 96.73%
inferior, or minimum diameter around the
conoid tubercle (99.63% dimorphism, with a female maximum exceeds the male) and the breadth of the
sternal facet (96.73%; also a case in which the female maximum exceeds the male). It is unsurprising, then,
that it is in these two dimensions that there are the smallest levels of dimorphism. The next smallest,
clavicle length, displays dimorphism of only 96.15% (p = 0.010) for which the mean values are significantly
different (male 129.29 mm, N= 19; female 124.31 mm, N=35). The anterior-posterior, or maximum,
diameter around the conoid tubercle is significantly dimorphic (86.64%, p = 0.001), as is the height of the
sternal facet (94.66%, p =0.037). It is interesting that at the sternal facet and the conoid tubercle, the larger
dimensions are significantly different, while the smaller ones are not. The maximum and minimum
diameters around the mid-shaft, however, are quite similar in their level of dimorphism (89.63%, p= 0.002
and 89.71%, p = 0.002) and are both significantly dimorphic. In addition, the ratio of the maximum mid-
shaft diameter to the clavicle length is sexually dimorphic 92.72% (p = 0.038). None of the other clavicular
dimensions, when divided by the length of the clavicle, demonstrated significant differences.
Table 13: Clavicular Model: discriminant function analysis
Eigenvalues Wilks' Lambda Standardized canonical discriminant Classification

function coefficients results
Eigenvalue % of Wilks' Chi- sq df sig male female

variance Lambda
0.573 100.00% 6.360 21.968 5 0.001 clavicle maximum length 0.468 77.80% 80.00%
clavicle mid-shaft diam (max) 0.468
clavicle mid-shaft diam (min) 0.503 Total: 78.90%
clavicle maximum (AP) diameter 0.271
at conoid tubercle
clavicle sternal facet (max) height -0.063
Sex identification based on the significantly dimorphic clavicular dimensions was biased in favour of
females, but had higher accuracy for both males and females than the model based on the scapular
dimensions. There was much less discrepancy between the contributions of each dimension to the overall
function than there in the scapular model. The minimum diameter around the mid-shaft of the clavicle
45
contributed slightly more to sex identification than did the other elements, although the sexual dimorphism
of the dimension is slightly less than that seen in the length of the clavicle, and it is statistically significant to
a similar level as the other dimensions.
Combining scapular and clavicular measurements results in a more effective model than either
skeletal element individually. There was no improvement from the clavicular model for female
identification, as 80% were correctly identified in both situations, but there was a vast improvement in
female identification from the scapular model. More importantly, using the entire shoulder girdle, males of
the sample are now identified correctly 93.8% of the time, an improvement from both previous situations.
The breadth of the scapula is still the most important dimension for sex identification, followed by the
scapula length as it had been in the first model; the importance of the clavicular dimensions shift slightly,
with the maximum mid-shaft diameter of the clavicle overtaking the minimum, and the clavicle length
dropping drastically.
Table 14: Shoulder Girdle Model: discriminant function analysis


variance Lambda
0.833 100.00% 0.546 26.962 9 0.001 scapula length 0.605 93.80% 80.00%
glenoid fossa length 0.010 Total: 86.90%
glenoid fossa breadth -0.105
clavicle maximum length -0.086

clavicle mid-shaft diam (min) 0.333
clavicle maximum (AP) 0.319
diameter at conoid tubercle
clavicle sternal facet (max) -0.072
height
46
Figure 5: Clavicle length, by sex Figure 6: Clavicle mid-shaft diameter (maximum),

by sex
Figure 7: Clavicle mid-shaft diameter (minimum),

by sex
Figure 8: Conoid tubercle diameter (anterior-

posterior), by sex
Figure 9: Clavicle sternal facet height (superior-

inferior)
47
Pelvis Table 15: Sex differences in the pelvis

Males are larger in all dimensions significance female mean as % of
level male mean
except for the superior pubic height, in os coxae height -- 97.83%
which the male mean (15.14 mm, N = 19) ilium breadth < 0.05 94.91%
Iliac crest (max) breadth -- 93.17%
exceeds the female (15.17 mm, N = 36) and
Iliac crest (min) breadth < 0.01 78.32%
the female maximum is larger than the acetabular height < 0.01 91.71%
male. This sex difference is the sole acetabulosciatic breadth -- 94.99%
superior pubic height -- 100.22%
instance in which the female mean is
symphyseal height -- 91.30%
absolutely larger than the male, with
symphyseal breadth < 0.05 86.18%
dimorphism of 100.22% (p = 0.950) but this auricular surface length 0.01 91.58%
is not statistically significant. Dimorphism is auricular surface breadth -- 91.43%

ratio of acetabular diameter to
< 0.01 93.85%
not statistically significant in the os coxae os coxae height
ratio of symphyseal breadth to
height (male mean = 276.06 mm, N = 18; os coxae height
< 0.5 87.77%
female mean = 270.06 mm, N = 35), the

maximum breadth of the iliac crest (male mean = 7.79 mm, N = 18; female mean = 7.26 mm, N = 35), the
acetabulosciatic breadth (male mean = 29.77 mm, N = 19; female mean = 28.27 mm, N = 36), the height of
the pubic symphysis (male mean = 51.86 mm, N = 19; female mean = 47.35 mm, N = 36), or the breadth of
the auricular surface (male mean = 15.31 mm, N = 19; female mean = 13.99 mm, N = 36).
The degree of dimorphism does not correlate to the significance; some of the dimensions which are
significantly different have levels of dimorphism within the range of those that are not significantly
different, including the breadth of the ilium (male mean = 124.74 mm, N = 18; female mean = 118.39 mm, N
= 35; 94.91%, p = 0.024) and the length of the auricular surface (male mean = 52.30 mm, N = 19; female
mean = 47.90 mm, N = 36; 91.58%, p = 0.009). The most obvious example of this is the comparison of the
length and breadth of the auricular surface, as the two have very similar levels of dimorphism (91.58% and
Table 16: Pelvic Model: discriminant function analysis

discriminant function coefficients results

variance Lambda
0.582 100.00% 0.632 22.261 5 0.000 ilium breadth -0.188 77.80% 80.00%
Iliac crest (min) breadth 0.766
acetabular height 0.423 Total: 78.90%
symphyseal breadth -0.032
auricular surface length 0.383
48
Figure 10: Ilium breadth, by sex
Figure 11: Iliac crest (minimum) breadth, by sex
Figure 12: Acetabulum height, by sex
Figure 13: Pubic symphysis breadth, by sex
Figure 14: Auricular surface length, by sex
49
91.43% respectively), yet it is only in the length that the sexes are significantly different. The minimum
breadth of the iliac crest (78.32%, p < 0.001) shows the highest amount of dimorphism, and with the height
of the acetabulum (91.71%, p< 0.001), is among the most strongly statistically significant sex difference. The
ratio of the height of the acetabulum to the height of the os coxae (93.85%, p = 0.005), and that of the
breadth of the pubic symphysis to the height of the os coxae (87.77%, p = 0.040).
Dimensions of the pelvis are only slightly better at identifying females, within this sample, than
males. The minimum breadth of the iliac crest, which is highly significantly dimorphic (p-value of < 0.01 )
and has the single greatest size difference of all pelvic dimensions, is the main contributor to this model.
The acetabular height follows in predictive value, despite having lagged behind other dimensions in size
difference; the remaining dimensions contribute minimally.
50
Forelimb
The length of the humerus Table 17: Sex differences in the humerus
(male 300.89 mm, N = 19; female significance female mean as % of
level male mean
mean = 298.94 mm, N = 36) varies humerus length -- 99.35%
very little between the sexes (99.35% humerus mid-shaft diam (max) < 0.05 95.17%
humerus mid-shaft diam (min) -- 95.07%
dimorphism) and is not statistically
humerus 35% diam (max) -- 95.88%
significant. The maximum diameter of humerus 35% diam (min) < 0.05 94.28%
the proximal head is significantly humerus proximal transverse diam < 0.01 92.23%
different between the sexes, although humerus proximal head diam < 0.01 92.17%
humerus surgical neck diam (max) < 0.01 88.48%
it varies to only a slight degree (male
humerus surgical neck diam (min) < 0.01 89.69%
mean = 41.71 mm, N = 19; 38.44 mm, humerus epicondylar breadth < 0.01 93.02%
N = 36; 92.17%, p < 0.001) as does the humerus distal articular breadth < 0.01 92.96%
ratio of humerus proximal diameter by
< 0.01 92.77%
transverse diameter of the proximal humerus length
ratio of humerus epicondylar breadth to
< 0.01 93.69%
epiphysis (male mean = 47.01 mm; N = humerus length
s
19; female mean = 43.36 mm, N = 36; 92.23%, p < 0.001). Both the minimum and maximum values of the
diameter of the surgical neck show significant levels of sexual dimorphism, and it is in these dimensions that
the absolute difference between the sexes' means shows the most dimorphism for any portion of the
humerus, with 88.48%, p < 0.001 (male mean = 28.47 mm, N = 19; female mean = 25.19 mm, N = 36) and
89.69%, p < 0.001 (male mean = 23.05 mm, N = 19; female mean = 20.67 mm, N = 36).
The diameter of the mid-shaft differs very slightly in its maximum value and minimum values,
although only the former is statistically significant, with females averaging 95.17% of the male size (male
mean = 24.89 mm, N = 19; female mean = 23.69 mm, N = 36). The opposite pattern is seen at 35% of the
way along the shaft: the slight variation in maximum and minimum values, the sex difference (94.28%, p =
0.019) of the minimum diameter (male mean = 22.19mm, N=19; female mean = 20.92mm, N=36) are
statistically significant. The distal articular breadth and the epicondylar breadth both differ slightly between
the sexes, but not to a significant degree.
Humeral dimensions can be used to predict sex within this sample with commensurate success for
both sexes. With 83.3% of males correctly identified and 85.7% of females, the bias in favour of females is
more slight than any other model based on a single bone. The epicondylar breadth is the most critical to
the model's power, followed by the minimum diameter around the surgical neck.
51
Table 18: Humeral Model: discriminant function analysis


variance Lambda
0.718 100.00% 0.582 25.437 8 0.001 humerus mid-shaft diam (max) -0.057 83.30% 85.70%
humerus 35% diam (min) -0.713
humerus proximal transverse 0.221 Total: 84.50%
diam
humerus proximal head diam 0.259
humerus surgical neck diam 0.401
(max)
(min)
humerus epicondylar breadth 0.723
humerus distal articular -0.382
breadth
Figure 15: Humerus proximal transverse diameter, Figure 16: Humerus proximal head diameter, by sex
by sex
52
Figure 17: Humerus surgical neck diameter Figure 18: Humerus surgical neck diameter
(maximum), by sex (minimum), by sex
Figure 19: Humerus mid-shaft diameter Figure 20: Humerus distal articular breadth, by sex
(maximum), by sex
Figure 21: Humerus epicondylar breadth, by sex Figure 22: Humerus 35% shaft diameter
(minimum), by sex
53
While the sex difference in humerus length

Table 19: Sex differences in the ulna and radius
was not significant, the lengths of the ulna and
level of male mean
radius were. There is more variation in the lower
ulna length (max) < 0.05 96.67%
arm than in the upper arm. The mean female ulna radius length (max) < 0.05 96.80%
length (mean = 285.08mm, N = 36) was 96.80% radius mid-shaft diam (max) < 0.01 92.20%
radius mid-shaft diam (min) < 0.05 94.18%
(p=0.025) of male (mean = 294.89mm, N = 19).
radius head diam < 0.01 92.26%
Female radius length (mean = 270.58mm, N = 36) a
was 96.67% (p = 0.023) the size of males (mean = 279.53mm, N = 19). Furthermore, all measurements on
the radius showed greater dimorphism than did the length: the diameter of the radial head (male mean =
25.86mm, N = 19; female mean = 23.85mm, N = 36) at 92.26% (p < 0.001) and the mid-shaft diameter
maximum (male mean = 16.23mm, N = 19; female mean = 14.97, N = 36) showed 92.20% (p = 0.007), while
the minimum diameter of the mid-shaft was slightly less significant, 92.26%, p= 0.017 (male mean =
13.80mm, N = 19; female mean = 13.00mm, N = 36).
Radial dimensions were, of all models using single bones, the least successful at identifying sex,
placing only 63.2% of males and 72.2% of females. The various dimensions contributed to the model's
success in decreasing order of the statistical significance of their sex differences.
Table 20: Radial Model: discriminant function analysis


variance Lambda
0.41 100.00% 0.712 17.352 4 0.002 radius length (max) 0.016 63.20% 72.20%
radius mid-shaft diam (max) 0.327
radius mid-shaft diam (min) 0.060 Total: 67.70%
radius head diam 0.817
54
Figure 23: Radius height, by sex
Figure 24: Radius mid-shaft diameter

(maximum), by sex
Figure 25: Radius mid-shaft diameter (minimum), by

sex Figure 26: Radius head diameter(maximum), by sex
55
Forelimb length (male mean = 580.42 mm, N = 19; female mean = 569.53 mm, N = 36) does not
differ significantly between the sexes. The brachial index, or the radial length as a percentage of the
humeral length, was not only significant, but was in fact the only of the limb indices to reach significance in
inter-sexual variation – this no doubt reflects the fact that the humerus lengths do not differ between the
sexes while the radial lengths do. Although the physical difference between the male mean (92.91 mm, n =
19) and the female (90.51mm, n = 36) is small, it is nonetheless highly significant with a p-value of < 0.001.
Combined humeral and radial dimensions do a better job of identifying sex than do the dimensions
of either element alone. It does not improve on female identification; the 85.70% of females identified in
both cases are therefore placed from humeral dimensions alone. The forelimb model has a 3%
improvement in male identification from the humeral model.
Table 21: Forelimb Model: discriminant function analysis


variance Lambda
0.755 100.00% 0.570 25.02 13 0.023 humerus mid-shaft diam (max) 0.009 88.30% 85.70%
humerus 35% diam (min) -0.803
humerus proximal transverse 0.246 Total: 87.00%
diam
humerus proximal head diam 0.276
(max)
(min)
humerus epicondylar breadth 0.679
humerus distal articular breadth -0.380
radius length (max) 0.205
radius mid-shaft diam (max) -0.128
radius mid-shaft diam (min) 0.288
radius head diam 0.016
56
Hindlimb
Femur length shows little, and
Table 22: Sex differences in the femur
statistically insignificant, dimorphism,
with the female mean a full 98.68% of level of male mean
femur length -- 98.68%
the male. Neither the minimum nor
femur mid-shaft diam (max) -- 94.41%
maximum values of the mid-shaft femur mid-shaft diam (min) -- 97.33%
diameter were significant, with 94.41% femur head diam (max) < 0.01 94.85%
femur SI height of the femoral neck -- 95.03%
and 97.33% dimorphism. While shaft
femur neck AP diam -- 95.81%
length and girth did not show sexual femur neck length (min) < 0.05 92.73%
dimorphism, nearly all of the epiphyseal femur biomechanical neck length < 0.01 92.14%
femur epicondylar breadth < 0.01 92.20%
measurements did; the only exception
femur distal anterior-posterior diam < 0.05 95.90%
being the the femoral neck, as both the ratio of femur biomechanical neck
< 0.01 93.13%
length to femur length
superior-inferior and anterior-posterior ratio of femur epicondylar breadth to
< 0.01 93.62%
femur length
diameters were insignificant in their
variation. Femoral head diameter, with the female mean (23.44mm, N = 19) 94.85% of the male (34.20mm,
N=36) was highly significant (p = 0.004). The other dimensions of the proximal epiphysis showed a similar
degree of dimorphism: the minimum length between the lesser trochanter and the femoral head 92.73% of
males (p = 0.034), and the biomechanical neck length (92.20%, p < 0.001).the epicondylar breadth 92.20%
(p < 0.001); the distal anterior-posterior diameter was slightly less dimorphic and slightly less significant, at
95.90% (p = 0.033). Additionally, the ratio of the femur biomechanical neck to the entire femur length (or,
the size-adjusted biomechanical neck length) was significantly dimorphic, at 93.13% (p < 0.001), as was the
ratio of the femur epicondylar breadth to entire femur length (93.62%, p < 0.001).
Femoral dimensions are relatively similar to humeral in terms of identifying sex; the model has less
accuracy for males but greater for females. The bio-mechanical neck length is by far the most contributive,
over and above the femoral head; expected, as it has the largest sex difference of those with the highest
statistical significance.
57
Figure 28: Femur biomechanical neck length, by sex

Figure 27: Femur head diameter (maximum), by sex
Figure 29: Femur distal diameter (anterior-

posterior), by sex Figure 30: Femur epicondylar breadth, by sex
Figure 31: Femur neck length (shortest distance

between head and lesser trochanter), by sex
58
Table 23: Femoral Model: discriminant function analysis


variance Lambda
1.067 100.00% 0.484 35.219 5 0.000 femur head diam (max) -0.601 84.20% 88.20%
femur neck length (min) 0.024
femur biomechanical neck 1.237 Total: 86.20%
length
femur epicondylar breadth 0.185
femur distal anterior- -0.042
posterior diam
The length of the tibia is not

Table 24: Sex differences in the tibia
significantly sexually dimorphic, but the
significance female mean as
diaphysis and joint surfaces were. Dimorphism level % of male mean
tibia length (max) -- 98.21%
of the mid-shaft in maximum diameter,
tibia mid-shaft (ap) diam (max) < 0.01 93.36%
females (mean = 23.73mm, N = 19) averaging tibia mid-shaft diam (min) < 0.05 94.40%
93.35% of males (mean = 22.15mm, N = 36), plateau breadth < 0.01 91.94%
ratio of tibia plateau breadth by tibia
was highly significant (p = 0.006); the 94.40% length
< 0.01 93.75%
dimorphism seen at the minimum diameter a
(male mean = 15.73mm, N = 19; female mean = 14.85mm, N = 36) was only just significant (p = 0.049). The
breadth of the plateau on the proximal epiphysis, with the female mean (55.38mm, n=19) 91.94%
(p<0.001) of the male (60.24, n = 36); the dimension was still sexually dimorphic when size adjusted,
although slightly less so, at 93.75% (p = 0.001).
The model to predict sex using dimensions of the tibia was somewhat efficacious, results biased in
favour of females at 86.1% and only 72.2% of males correctly placed. Plateau breadth far surpassed mid-
shaft dimensions in contribution, implying that despite comparable degrees of significance, the differences
in joint surface may be more important than those the diaphysis.
Table 25: Tibial Model: discriminant function analysis

discriminant function coefficients results

variance Lambda
0.641 100.00% 0.609 25.020 3 0.000 tibia mid-shaft (ap) diam 0.172 72.20% 86.10%
(max)
tibia mid-shaft diam (min) 0.054 Total: 79.15%
plateau breadth 0.922
59
Figure 32: Tbia mid-shaft diameter (maximum), by Figure 33: Tibia mid-shaft diameter (minimum), by sex
sex
Figure 34: Tibia plateau breadth, by sex
60
Hind-limb length (male mean = 549.67mm, N = 18 and female mean = 541.00mm, N = 36),
calculated simply as the sum of the femur and tibia lengths, did not vary significantly between the sexes;
the lack of significant difference in the femur must have overshadowed that of tibia length. Neither the
intermembral index nor the crural index showed a significant sex difference.
In terms of predictive ability, the combination of femoral and tibial dimensions had greater accuracy
than had either element individually, as expected. A full 94.1% of females were placed and 88.9% of males.
This success rate surpassed not only the the model based on forelimb dimensions, but even that of the
shoulder girdle. Of any region of the body, the hindlimbs seem to be the strongest predictor indicative of
sex, at least in this model within this sample.
Table 26: Hindlimb Model: discriminant function analysis

discriminant function results
coefficients
variance Lambda
1.532 100 0.4 42.726 8 0.000 femur head diam -0.751 88.90% 94.10%
(max)
femur neck length 0.081 Total: 91.50%
(min)
femur biomechanical 1.135
neck length
femur epicondylar -0.194
breadth
femur distal anterior- -0.336
posterior diam
tibia mid-shaft (ap) 0.186
diam (max)
tibia mid-shaft diam 0.075
(min)
plateau breadth 0.783
61
Body proportions
Presented in Table 9 are the biometric data taken
from Merfield's field notes. Means and standard deviations
were calculated for males, females, and a combined sex
total to represent the population as a whole. For none of
the attributes measured were the differences between the
means by sex statistically significant; that is not to say that
these features do not demonstrate sex differences, but that
Figure 35: “Belly” measurements for a
they are not apparent in the available data – probably due subset of the sample, from 1933 field notes
to the small sample sizes. The fact that in a few cases (body
height, belly, and hand height) the female mean exceeds
the male mean is interesting, but given the lack of statistical
significance, it is too early to draw any firm conclusions. It is
entirely feasible that females have a larger circumference
around the “belly”, particularly if the sample included any
pregnant females (although no pregnancies are mentioned
in the notes, the possibility cannot be discounted). There is
Figure 36: “Bicep girth” measurements for
less reason to consider the suggestion that the average a subset of the sample, from 1933 field
female height is taller than that of males; the average notes
female femur length, tibia length, or os coxae heights are all

smaller than the male averages, but as the differences are
not statistically significant, it is impossible to conclude that
the height difference in the field notes is inaccurate or
merely a distortion from an inadequate sample size.
Figure 37: “Chest” circumference

measurements for a subset of the sample,
from 1933 field notes
62
Figure 39: “Forearm” length

measurements for a subset of the sample, Figure 38: “Girth” measurements for a
from 1933 field notes subset of the sample, from 1933 field notes
Figure 40: “Hand” height measurements Figure 41: “Height” measurements for a
for a subset of the sample, from 1933 field subset of the sample, from 1933 field notes
notes
Figure 42: “Neck” circumference Figure 43: “Span” measurements for a

measurements for a subset of the sample, subset of the sample, from 1933 field notes
from 1933 field notes
63
the skeletal measurements, indices and ratios were calculated. Table 7 presents the basic statistics, and
independent sample t-tests comparing the female and male means. In addition to the brachial, crural, and
intermembral indices which consider long bone lengths, several additional indices were added that involved
the clavicle, as a stand-in for torso girth. With the exception of the brachial index, none of the indices
comparing different bones demonstrated significant sex differences. Although none of the indices
comparing the clavicle to long bones reached statistical significance, claviculo-humeral index was the
closest, with a p-value of 0.06.
To determine the correlation between trunk dimensions and sex identification, a model was used
that combined the available significant clavicular, scapular, and pelvic dimensions. Dimensions from each of
the bones contributed in relatively similar degrees, although the greatest contributions were made by two
scapular measurements. It was quite successful at predicting the sex of specimens in the sample, correctly
placing 87.5% of the males and 88.2% of females. It was an improvement overall from the model using only
the shoulder girdle, which while able to place 93.8% of males, correctly identified 86.5% of the cases,
whereas the trunk model placed 87.85% overall. The difference between these success rates is so slight that
without comparing to other populations, or dealing with a larger sample, it is unclear as to whether this is a
true improvement.
Finally, the proportional calculations were used in a last discriminant function analysis to predict
Table 27: Trunk Model: discriminant function analysis


variance Lambda
1.168 100.00% 0.461 31.726 14 0.004 scapula length 0.593 87.50% 88.20%
glenoid fossa length -0.144 Total: 87.85%
glenoid fossa breadth -0.268
clavicle maximum length -0.082
clavicle mid-shaft diam (min) 0.298
clavicle maximum (AP) 0.371
diameter at conoid tubercle
clavicle sternal facet (max) -0.127
height
ilium breadth -0.417
Iliac crest (min) breadth 0.554
acetabular height 0.007
symphyseal breadth -0.090
auricular surface length 0.338
64
sex. The brachial index was combined with measurements divided by the lengths of the elements from
which they came, i.e. size-adjusted values. This model is successful with rates comparable to others, as it
identified 85.7% and 84.80% of females. While some of the other models have higher success rates, this
model is perhaps more interesting; the fact that it is still successful at identifying sex without using absolute
size values means that there is not only a pattern of proportional shape differences between males and
females, but that these shape differences are strong enough to identify sex. It would be valuable to
compare this to data from other populations.
Table 28: Proportional Model: discriminant function analysis


variance Lambda
1.001 100.00% 0.500 28.087 9 0.001 brachial index 0.644 85.70% 84.80%
clavicle mid-shafter diameter 0.138
(max) to clavicle length
acetabular diameter to os 0.113 Total: 85.25%
coxae height
symphyseal breadth to os 0.284
coxae height
humerus proximal diameter by -0.040
humerus length
humerus epicondylar breadth to -0.064
humerus length
femur biomechanical neck 0.554
length
femur epicondylar breadth to 0.102
femur length
tibia plateau breadth by tibia 0.217
length
65
Discussion and Conclusions

The results of this analysis confirm the complicated, mosaic nature of sexual dimorphism in
chimpanzees. Many of the dimensions found to have significant sexual dimorphism cannot be compared to
the literature as this study, in the attempt to provide more thorough and rich data, includes dimensions that
simply have not been previously addressed. Additionally, a number of measurements that were not found
to be significant in previous work, but are in this study, likely reflect differences in this population relative to
others; however, that this analysis is based on a much larger sample size may contribute to that impression.
Measured values, comparison to previous studies

The results of this study are in line with previous studies. See Table 1 for available information from
previous publications. Males are absolutely larger, with the exception of measurements of the pubis. Means
are within a similar range to those previously recorded, suggesting that these measurements are indeed
accurate; the fact that the majority of measurements demonstrate larger averages, for both males and
females, indicates by its consistency that the population from which the sample was taken is, in entire body
mass, larger than those to which it is compared. This is unsurprising, as intra-specific size variation is well
known in Pan troglodytes. Perhaps the strongest piece of evidence which supports the accuracy of these
measurements is that the indices, or relations of different skeletal elements to each other, fall even closer
to previously published values than do the measurements themselves. The degree of sexual dimorphism
measured does vary from dimension to dimension, relative to previous studies, but this does not
undermine the results; previous work has indicated that sexual dimorphism varies between subspecies and
populations, both in absolute degree and as a matter of which specific dimensions demonstrate statistically
significant differences.
Dimensions of and sex differences in the shoulder girdle

Zihlman et al (2008) found female mean scapula breadth to be 87.8% of males; this analysis of
chimpanzees from Cameroon found slightly less sexual dimorphism in the same measurement, with an
average of 94.35%. The Tai population has a male mean of 80.2 mm (N=5) and female 70.2 mm (N = 10), as
compared to 86.85 mm (N = 17) and 81.94 (N = 36) in this sample – meaning, the difference between the
sexes is comparable, but the Tai chimps of both sexes were smaller. Zihlman compared the Tai population to
the Pan troglodytes schweinfurthii of Gombe, whose measurements fell between the Tai and those
reported here, with a male mean scapula length of 83.4 mm (N = 3) and a female mean of 73.0 (N = 8),
resulting in 87.5% dimorphism. Clavicle lengths are reported for the Tai and Gombe populations, with 92%
66
dimorphism at Tai and 91.7% at Gombe (although this difference was not statistically significant). This
analysis again finds slightly less, with dimorphism at 96.15%. The mean values (Tai males = 121.3 mm, N = 5,
females = 111.5 mm N = 9; Gombe males = 116.6 mm, N = 5 and females = 107 mm, N = 8) are once again
smaller than those in this population (males = 129.29, N = 19; females = 124.31, N = 35).
Dimensions of and sex differences in the pelvis

Dimorphism of the os coxae length was not significant in this population, nor had it been at Tai or
Gombe. This Cameroonian population shows 97.83% dimorphism (combined sex average = 272.09, N = 53),
compared to 96.8% (combined sex average = 267.6, N = 13) in Tai and 95.9% (combined sex average = 258.2,
N = 13) at Gombe. Morbeck and Zihlman (1992) report a combined sex average for Gombe chimps of 254.9
mm (N = 9). Dimorphism in this population is similar, with slightly less found in this population and absolute
values slightly larger. Published values for dimorphism in the breadth of the ilium varies from 91% (at
Gombe in Morbeck et al 1992) to 93.2% (Gombe, in Zihlman et al 2008). The degree found in this sample is
less, with the female average a full 94.91% of males, but unlike previous studies, it is statistically significant.
With dimorphism of 104% as reported by Morbeck et al (1992), pubis height is one of the few
measurements favouring females as the larger sex. Although I did not measure pubis length in this sample,
it can be roughly compared to the superior pubic height, which at 100.2% dimorphism also favours females.
The diameter of the acetabulum was not significant in previous studies, although it is in this
analysis: the sex difference is highly significant (p-value < 0.001) with the male mean = 39.57 mm , N = 19
and the female mean = 36.29 mm, N = 36. The 94.99% degree of dimorphism is the first instance to this
point in which this analysis finds a greater amount of dimorphism, rather than less, from previously
published values; the dimorphism of the acetabular height, is intermediate between the 96.7% of the Tai
population (male mean = 39.1 mm, N = 6; female = 37.8 mm, N = 8) and 94.5% at Gombe (male = 35.9 mm,
N = 5; female = 34.1 mm, N = 8) as described by Zihlman et al (2008). Morbeck et al (1992) reported an 11%
size difference in a Gombe sample.
Dimensions of and sex differences in long bones

Of the long bones, the only measurements available from previous studies for comparison are the
lengths. Humerus length in this sample averages 299.62 mm (N = 55) for the sexes combined, 300.89 mm (N
= 19) for males, and 298.94 mm (N = 36) for females. These values are close to those of previous studies
and only slightly greater: at Tai, the male average was 297.9 mm (N = 5) and the female average was 296.8
mm (N = 9); at Gombe, the male average was 277 mm (N = 6) and the female 271 mm (N = 8). The sex
difference in these three (99.35%, 99.60%, 98.10% respectively) are all less than the 89.00% published in
67
Morbeck et al (1992), but close to the 97.71% dimorphism described by Schultz (1937), derived from a male
average of 306 mm (N = 45) and female average of 299 mm (N = 62). The combined sex average from
Morbeck & Zihlman (1989) of 301.6 mm (N = 14) is higher than any other, including that found at this time
for the Cameroon sample; this is intriguing, as it deviates from the pattern in which this study has so far
found larger values than were known previously. The relative mean lengths of the radius are similarly
distributed: Zihlman et al (2008) found a male average of 272.1 mm (N = 4) and a female average of 268.9
(N = 9), resulting in 98.8% dimorphism, at Tai and 96.6% at Gombe, from averages of 256.8 mm (N = 6) for
males and 246.3 mm (N = 8) for females. The findings here demonstrate a degree of dimorphism similar to
Gombe, although larger dimensions than either site, as males average 279.53 mm (N = 19), and females
270.58 mm (N = 36 ) for 96.8%. Femur lengths reported by Zihlman et al (2008) for the sample from Gombe
(male = 274.3 mm, N = 5; female = 246.3 mm, N = 7) are the smallest, with Tai (male = 297 mm, N = 4;
female = 288.9 mm, N = 9) demonstrating larger values, but the two samples have a similar degree of
dimorphism (in this case, exactly the same, 97.3%). In this sample, males average 296.56 mm (N = 18) and
females 292.64 mm (N = 36), larger than either the Tai or Gombe samples, but demonstrating less
dimorphism (only 98.68%). Tibia length follows the now familiar pattern: Gombe values are the smallest but
have the most dimorphism, values at Tai are intermediate in size, and those here the largest. Tai (male =
243.3 mm, N = 6; female = 239.6 mm, N = 7) has 98.5% dimorphism, Gombe (male = 230.8 mm, N = 5;
female = 222.7 mm, N = 7) has 96.50%, and in this study at Cameroon (male = 252.89 mm, N = 19; female =
248.36 mm, N = 36) has 98.21%.
Despite the differences in average measurements, ratios between elements are quite similar. The
intermembral index, for combined sexes, is 106.7 at Tai and 105.9 at Gombe according to Zihlman et al
(2008). Schultz (1937) reported average by sex, with 107.3 (N = 44) for males and 106.4 (N = 58) for females.
This is similar to that in this sample, 105.74 for males and 105.35 for females. The humeral-femoral index at
Tai averages 102.2, and at Gombe of 101.4; in this study, the indices by sex straddle those values, with
101.62 (N = 18) for males and 102.23 (N = 36) for females. The crural indices fall within a small range: 81.9
at Tai, 83.8 at Gombe, in Schultz (1937), 83.5 (N = 45) for males and 83.4 (N = 60) for females, and in this
sample, 85.37 (N = 18) for males and 84.87 (N = 36) for females. Schultz (1937) reported claviculo-humeral
index averages of 42 (N = 35) for males and 42.2 (N = 52) for females; this is very similar to those found in
this analysis (male = 43, N = 18; female = 41.69, N = 35). The brachial index has combined sex averages of 90
at Tai, 91.8 at Gombe, and 91 as in Morbeck & Zihlman (1989). In this analysis, it demonstrated a highly
significant sex difference as male = 92.91 (N = 19) and female = 90.51 (N = 36); this is similar to those from
Schultz (1937), a male average of 93.1 (N = 45) and female average of 92.6 (N = 60).
68
Trunk size
The scapulas of males and females differed in length and breadth – the breadth is more dimorphic
than the length (both in absolute value and in statistical significance) and may reflect males' broader, wider
trunks. Numerous muscles attach to the scapula, which can be said to “hang” in an envelope of muscle. I
had expected to find more of the scapular dimensions significantly sexually dimorphic; several of the
dimensions measured were added midway through the period of data collection, to focus on areas that
appeared highly variable (specifically: spine length, spine height, coracoid length, acromion breadth, or
superior border length). The acromion process articulates with the clavicle in the acromioclavicular joint;
the coracoid process meets ligaments which attach to the acromion process and the clavicula, to stabilize
the acromioclavicular joint, as well as the three muscles responsible for the movement of the arm ; and the
scapular spine serves as muscle attachments in addition to stiffening the entire bone. While none of the
additional dimensions showed significance in sex difference, several were, in fact, more variable than other
scapular dimensions: the length of the superior border, the height of the spine, and the acromion breadth
had coefficients of variation (0.2050, 0.1158, and 0.1587 respectively) that were well above all other
dimensions, which ranged from 0.0561 to 0.0726. In all of these cases, females were more variable than
males (as demonstrated by higher CVs); as this was also true for the sexually dimorphic dimensions, females
were more variable in every scapular dimension measured with the exception of the coracoid length. The
question of how to explain this variation, at an individual level as opposed to by sex, is intriguing. Given that
these dimensions relate to areas of muscle attachment and that an individual's musculature is related to
their nutritional status, it seems relevant to point out that young adult chimpanzees of both sexes are
smaller than more mature and higher-ranking individuals: body mass dimorphism peaked at 13 years of age
(Pusey et al 2005), but the maximum body mass measured for either sex was closer to 25 years of age
(Goodall 1986)and, additionally, female body mass is highly correlated with social rank (Pusey et al 2005). It
is possible that the non-sex variation in scapular dimensions reflects age or rank-related variation in body
mass.
Significant sex differences in the clavicle range from 86.64% to 96.15%. The amount of variability
within the combined-sex sample of the non-significant (p-values > 0.05) dimensions is within the range of
variability of those which were significant. Despite the fact that clavicle length showed the smallest amount
of overall variation (CV = 0.0541, while all dimensions ranged from 0.0928 to 0.1584), it can be considered
a strong predictor of sex, having been significantly dimorphic in both this analysis and in the Tai population
(Zihlman et al 2008). The clavicle is critical to mobility and function of not only the shoulder and forelimb
girdle, but the entire primate trunk. In mammals that have lost the clavicle, the upper limb girdle is attached
to the trunk solely by musculature; in primates, the clavicle acts as a strut to keep the scapulae apart and in
69
a lateral position, allowing a higher degree of mobility in the articulation of the scapula and humerus.
(Ankel-Simons 2007) The fact that it is the last bone in the body to fuse (White & Folkens 2005, 195) may
allow for a higher degree of variability than would be expected given the critical nature of its functional
purposes. Generally speaking, primates with more barrel-shaped torsos have longer clavicles than do those
with longer, more narrow trunks. As male chimpanzees have broader torsos than do females, a hypothesis
would follow that the clavicles of males are longer, relative to their other dimensions. (Ankel-Simons 2007)
To test this, proportions of the various dimensions of the clavicle were calculated relative to clavicular
length. Results, while failing to confirm the hypothesis, are somewhat equivocal: the ratio of the maximum
mid-shaft diameter to clavicle length does demonstrate a significant sex difference (p-value = 0.038, male
mean ratio of the two measurements is 0.1026, N = 18; female mean = 0.0951, N=36), but the larger ratio of
the males means that their clavicles are actually shorter relative to their diameter. Given that male clavicles
are indeed longer overall, that all other dimensions when calculated as ratios failed to reach significance,
and the likelihood that functional constraints on clavicular dimensions prevent a large amount of individual
variation, the hypothesis cannot be entirely dismissed.
The pelvis surrounds the birth canal in females and is subject to stress from locomotor activities and
postural behaviour. Comparison of the human pelvis to those of extant apes reveals not only the functional
adaptations which bipedalism has required, but also a variety of weight-bearing constraints. The human
pelvis is highly sexually dimorphic, but the majority of the features which distinguish the sexes involve the
maximization of the birth canal in females. As chimpanzee infants are relatively smaller at birth, the
anatomical structure of the female pelvis is less confined to parturition. In non-human primates there is an
allometric relationship between the size of the head of the foetus and the size of the mother's birth canal;
smaller species tend to give birth to infants with relatively larger heads. The pelvis is formed by two os
coxae, each of which consists of three bones which have permanently fused together. The hip bones meet
in two places: dorso-caudally at the sacral articulation and ventro-caudually at the symphysis. The auricular
surface, or the sacroiliac joint where the sacrum articulates with the os coxae, is the most immobile synovial
joint in the body. (White and Folkens 2005). The pelvic symphysis fuses late in life, although it may loosen
during the birth process in females: Morbeck et l (1992) concluded that this was not the case in
chimpanzees, due to the lack of “distinctive resorption areas” on the dorsomedial pubis and preauricular
area of the ilium, in agreement with Tague (1988). The lack of resorption areas in female chimps would
challenge the understanding that they are a necessary result of parturition. The dimorphism that this
analysis found breadth of the pubic symphysis and the length of the auricular surface seem to be
unprecedented in previous work, but implies that there may be more variation in those regions than has
been assumed. The iliac crest was dimorphic in its minimum width, although not in its maximum. This raises
70
the possibility that the maxima width is constrained due to relating more directly to muscular articulation
and locomotion, while the minima varies with body mass and morphological robusticity.
Joints
At its distal epiphysis of the humerus, the gleno-humeral articulation joins the humerus to the
scapula. The length and breadth of the glenoid fossa are significantly dimorphic (95.56% and 95.12%) as
was the maximum diameter of the humeral head (92.17%). While the existence of a sex difference in the
shoulder joint might initially seem counter-intuitive (the importance of the shoulder in movement and
locomotion specific to the species implies a high degree of functional constraint), that the difference in
trunk size between the sexes involves a concomitant difference in the shoulder joint, which supports the
trunk, is not inconsistent with previous data which demonstrates locomotory differences in male and
female chimpanzees, including in their suspensory behaviours. That other dimensions of the proximal
epiphyses also demonstrated significant sex differences further supports this conclusion.
Measurements of the distal end of the humerus, dealing with the elbow joint, demonstrate strong
sex differences which are, as these measurements have not previously been analysed in this manner,
unprecedented in studies dealing with chimpanzee variation. The epicondylar breadth of the humerus, the
distal articular breadth, and the maximum diameter across the radial head show dimorphism between 92%
and 93%.These dimensions are also highly critical to movement and suspensory behaviour. The lateral
trochlear ridge of the humerus is particularly well-developed in apes as it stabilizes the articulation between
the ulna and the distal humerus throughout forelimb extension and flexion, with olecranon fossa stabilizing
the joint in hyper-extension, which is known to be extremely important in knuckle-walking (Aiello & Dean
2005). That such a feature might differ between male and female chimpanzees must relate to the greater
amount of time that male chimpanzees spend in terrestrial locomotion (Doran 1993). The sex difference
displayed in the brachial index for this sample would support this interpretation. In addition, the
epicondylar breadth relative to humerus length is also significantly dimorphic for this sample, meaning that
the differences are not limited to size. A closer and more detailed consideration of this region would
provide a more detailed picture, perhaps increasing the amount of sex difference expected.
The acetabulum, or articular fossa, holds the head of the femur; the fossa itself is where the three
bones of the os coxae meet and have fused together. Cartilage forms the lunate articular surface within the
acetabulum, which does not extend across the entire perimeter of the acetabulum, but is open at the
acetabular notch; the wider the articular band is, the deeper the acetabulum. In general, the deeper the
acetabulum the more restricted the movements of the hip joint. The acentabulum of humans is
considerably more shallow than those of other primates and African apes (although deeper than
71
orangutans). Whether those of chimpanzees are deeper than humans is unclear: Schultz (1969b in Aiello
and Dean 2002) stated that they did, but Ruff (1988) found the opposite. Aiello and Dean (2002) note that
the high amount of intraspecific variation in acetabular depth may play a role in complicating such a cross-
species comparison. The acetabulum height does not, within this sample, seem to be more variable than
other pelvic dimensions: the coefficient of variation for acetabulum height (0.1190), while that for os coxae
height (a dimension which is not dimorphic) is 0.0440, and symphyseal breadth, highly dimorphic, is 0.2283.
While this indicates that the dimension is no more variable between individuals within this population, it
does not preclude the possibility that it is highly variable between populations. The height of the
acetabulum did show a strong sex difference in absolute size, as well as when considered as a ratio against
the height of the entire os coxae – meaning the acetabulum displays sex differences both absolutely and
proportionately. Dimorphism does not seem to vary dramatically between this sample, Tai, and Gombe.
Both the amount of dimorphism and the average values themselves are very similar in this sample and that
of Tai, but this may not be a fair comparison as the two may share a subspecies affinity.
In addition to the femoral head diameter – in which females averaged 94.85% of the male size –
other proximal dimensions of the femur were dimorphic, including the minimum length between the lesser
trochanter and the femoral head and the length of the biomechanical neck (although both superior-inferior
and anterior-posterior diameters of the femoral neck were not sexually dimorphic). With averages of
92.73% and 92.14% and comparable significance levels, there is actually more dimorphism in these
measurements than in the femoral head or the acetabulum diameter. The other measurements, more
commonly used throughout palaeoanthropological and primatological interpretations, may provide under-
estimations of the degree of musculature-related sex difference in the hip joint. The dimorphism of
measurements taken at the distal epiphyses of the femur and the proximal epiphyses of the tibia were
highly significant, with size differences of 92% to 96%. There are only three muscles which act exclusively on
the knee joint, and the femoral epicondyles provide a site of attachment for ligaments which stabilize it.
Long bone: lengths and diaphyseal properties

The sex differences in the lengths of long bones has already been addressed: humerus, femur, and
tibia length did not differ significantly between the sexes, but the average ulna and radius lengths did
display significant dimorphism, and neither total forelimb nor total hindlimb length demonstrated
significant difference. The obvious implication is that forelimbs are more variable than the upper arm (and
the lower leg more so than the thigh), although the coefficients of variation for the humerus, ulna, radius,
femur, and tibia were remarkably similar (all falling between 0.0431 and 0.0546).
While the epiphyseal dimensions of the long bones, forming joint surfaces, have historically
72
received more attention than the bone shafts, shaft dimensions and robusticity are important when
considering variation, both intra-specific and cross-species, as they reflect both phylogenetics and life
history variables, including development and habitual behaviour (Stock & Pfeiffer 2001; Stock 2001; Stock
2006). Robusticity was originally used to refer to the diaphyseal thickness standardized to bone length
(Stock & Shaw 2007), but more sophisticated methods of quantifying skeletal and morphological
characteristics have been developed (notably, cross-sectional geometry) that directly consider the
mechanical properties.
In the humerus, sex differences in diaphyseal diameter were apparent at both the 35% (in minimum
diameter) and 50% (in maximum) points, but were slighter than the dimorphism seen in the epiphyseal
dimensions. Radius mid-shaft diameter was significant in both maximum and minimum, femur shaft
diameter was dimorphic in its maximum , and the tibia mid-shaft significantly different in both maximum
and minimum diameters.
Sex identification
Discriminant function analysis classified the sex of specimens based on subsets of their
measurements: those of scapular dimensions, clavicular dimensions, the shoulder girdle (scapular and
clavicular combined), pelvic dimensions, trunk dimensions (scapular, clavicular, and pelvic combined),
humeral dimensions, radial dimensions, forelimb dimensions (humeral and radial combined), femoral
dimensions, tibial dimensions, hindlimb dimensions (femoral and tibial combined). Rather than using the
entire dataset for sex identification – which would no doubt have resulted in a higher accuracy rate –
subsets were chosen, so as to demonstrate the relative strength of sex differences in various anatomical
regions of the body, and to create a more realistic situation that may arise in field conditions and future
research. The fact that high levels of accuracy can be achieved using single elements or anatomical regions
attests to the significance in the pattern of sexual dimorphism, and has important implications for
palaeoanthropological and archaeological studies (in which only single elements are often available).
Correct classification rate of more than 80% were achieved by all but two models (radial dimensions had
67.7% accuracy, and tibial 79.15%). Models using dimensions of single skeletal element had accuracy rates
of more than 85% in two cases: humerus (86.2% based on femoral dimensions and 84.5% based on
humeral).
The greatest successes in identification were achieved with models combining measurements from
related skeletal elements: the dimensions of the trunk, reached 87.85% accuracy and dimensions of the
hindlimb reached 91.5%. While these rates are impressive – particularly given that they are based entirely
on postcranial dimensions, while traditional methods of sexing chimpanzee skeletal remains refer to cranial
73
and dental features – and are within the range of what researchers consider successful for humans, it will
be interesting to see if they are similar in other populations and other samples. If sexual dimorphism in
chimpanzees is as mosaic in its intra-specific variation as is currently thought, it is likely that the relative
accuracy of these models may reverse their relationships.
Many skeletal dimensions overlap in size between males and females, as does body mass.
Proportional relationships between dimensions, if found to demonstrate significant and strong patterns of
sex differences, are preferable for use in sex identification as they would preclude confusion with
individuals who are unusually large (or small) for their sex. The discriminant function analysis based only on
size-adjusted, or proportional, calculations placed 85.25% of specimens correctly. Despite being slightly less
than the former two models, its results are substantial and of great interest; clearly, sexual dimorphism in
chimpanzees dimorphism is not allometric, and not entirely a function of size.
Implications for intra-specific variation

This analysis is done with the assumption that the chimpanzees of the Powell-Cotton collection are
of the same subspecies; it is unclear, given the range of localities and the fact that they were collected at
various times, whether they represent a biological population. The Powell-Cotton collection has, in many
previous studies, been analysed collectively. It has seemed fair to assume that, whether or not they formed
a living population, they share a greater affinity with each other than to specimens from other sites or time
periods. It is, however, worth noting that actual taxonomic affinity of the collection may be debatable: not
only could the sample represents Pan trolodytes troglodytes, or Pan troglodytes verus, the “new”
subspecies Pan troglodytes vellerosus (or Pan troglodytes ellioti) is a candidate as well, given that it is
native to Cameroon. Alternatively, the collection could represent multiple subspecies. The field notes from
the expeditions include detailed descriptions of pelage colouration and body types, with frequent mention
of the local guides' names and descriptions of the specimens they had shot, as well as outright conjecture
as to the specimens' classification. The author is clearly under the impression that he is familiar with, and in
the process of collecting, specimens of at least two different types.
Assuming that the sample is, in fact, taxonomically heterogeneous would in no way invalidate the
results or interpretations herein; it would, however, alter the implications they have on the amount and
nature of intra-specific variation in Pan troglodytes' sexual dimorphism. As it seems that sex differences vary
between populations (with some populations displaying a greater absolute degree of dimorphism, perhaps
in different skeletal elements or different dimensions), the heterogeny of a sample would presumably
reduce the number of measurements in which dimorphism was found to be statistically significant. On the
other hand, if there is a size variation between populations (regardless of whether they share the same
74
pattern of sex differences) the admixture could result in an overestimation of the degree of dimorphism of
individual features. When this study is compared to previous datasets, it seems that neither of those
projections are realized: the amount of dimorphism in this sample is less, in nearly all dimensions
measured, than was seen in a sample of Pan troglodytes verus from Tai – and there is an increase in the
number of dimensions that display significance.
Whether these “newly” dimorphic dimensions, significant in this sample but not in previous, are an
artefact of the increased sample size or represent a taxonomic difference (and if so, at what grade:
population or subspecies?) is a question that deserves future attention. Further alternatives include the
possibility that the differences are a function of ecological changes (and if so, through what mechanism:
genetic or developmental?). Of course, these two hypotheses are not mutually exclusive. Regardless of
precise taxonomy, the chimpanzees of this sample from Cameroon would be more closely related to those
of Tai than the other historically well-studied populations (ie, Gombe and Mahale in Tanzania). However,
this sample does pre-date Tai by several decades. Secular change in chimpanzee skeletal material has not
been thoroughly examined yet, however Morbeck (1999) found preliminary evidence for such at Gombe.
The nature of those changes is, however, less clear. The simplest environmental change about which to
hypothesize is nutritional deprivation; this is hardly a hypothetical problem for great apes, given the drastic
habitat loss that they have endured in the last century and the fact that they are currently on the verge of
extinction. That wild chimpanzees at any site are undergoing habitat restriction and, therefore, relative loss
of dietary resources would seem to be a justifiable premise from which to operate. If nutritional deprivation
in chimpanzees has the same effect on growth as it does in humans (eg, males suffer more drastically with
higher rates of stunting, thereby decreasing the appearance of dimorphism across the population), we
could hypothesize that this sample, being an earlier population, was under less nutritional stress, and would
therefore display more sexual dimorphism – or, that descendent populations, under more stress, will be
displaying less sexual dimorphism as a result of males getting smaller. In almost all dimensions measured,
this sample is larger than Tai for combined sex total: these males from Cameroon are indeed larger than
males from Tai (or Gombe) – however, the females are larger as well (and in many cases, by a larger
margin). More importantly, the percentage of size difference in nearly all dimensions is less for the
Cameroonian sample (as opposed to more). All the same, as these comparisons are made simply from
statistical summaries, it may be premature to dismiss such a hypothesis without a more detailed intra-site
analysis. Secular and ecological changes – how they may affect sex differences, and the ways in which they
might be read in the skeletal record – are an exciting direction for further study that, hopefully, detailed
analyses such as this one may enable.
75
Chimp sexual dimorphism is not fully understood. Even human sexual dimorphism can only be
appraised within population context, and we have not yet built a sufficient framework to do the same with
chimpanzees. Nonetheless, palaeoanthropology has relied on data from chimpanzees for reconstruction, for
sex identification, and from which to draw behavioural hypothesis. Palaeoanthropological considerations of
sexual dimorphism focus on long bones (particularly femur), but previous studies demonstrate that this may
not be a good indicator of sexual dimorphism and there is, therefore, a need to evaluate the ability of
estimating from other elements. The current understanding of chimpanzee skeletal anatomy suggests that,
with the exception of canines, sex differences are minimal. Morphological comparisons between
populations and subspecies have shown that differences are due to female variation. Sex differences seem
to present as a mosaic, with different traits varying to different degrees in different populations. The results
of this study, however – demonstrating the substantial sexual dimorphism throughout the entire body,
across features which serve various mechanical functions – challenge these assertions.
76
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Postcranial Sexual Dimorphism in Pan Troglodytes - UOC THESIS 2012 NOELLE TANKARD

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Postcranial Sexual Dimorphism in Pan Troglodytes - UOC THESIS 2012 NOELLE TANKARD

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A dissertation submitted in partial fulfilment of the requirement for a

Master of Philosophy in Human Evolutionary Studies

Supervisor: Jay Stock

Postcranial Sexual Dimorphism

Sexual dimorphism in primates

and nature as well as in degree (Plavcan 2011).

Human sexual dimorphism

Plasticity and development

Quantifying sexual dimorphism

Geographic variation in chimpanzees and subspecies

Body size: dimorphism and proportions

Literature Review: Previous studies of postcranial sexual

Zihlman et al 2008 Zihlman1989

Table 2: Measurements of the scapula

Table 3: Measurements of the clavicle

Table 4: Measurements of the pelvis

Table 5: Measurements of the humerus, ulna, and radius

Table 6: Measurements of the femur and tibia

Table 7: Limb lengths, ratios, and indices

Table 8: Sexual dimorphism in proportions

Table 9: Biometric data from field notes, Merfield 1933

Summary statistics Independent samples t-test

ranging from 98.16% dimorphic (coracoid length) acromion breadth -- 93.78%

Table 11: Scapular Model: discriminant function analysis

Eigenvalues Wilks' Lambda Standardized canonical Classification

Figure 1: Scapula lengths by sex

Figure 3: Glenoid fossa length Figure 4: Glenoid fossa breadth

In all measured dimensions of the

Table 13: Clavicular Model: discriminant function analysis

Eigenvalues Wilks' Lambda Standardized canonical discriminant Classification

Eigenvalue % of Wilks' Chi- sq df sig male female

Table 14: Shoulder Girdle Model: discriminant function analysis

Eigenvalues Wilks' Lambda Standardized canonical discriminant Classification

Eigenvalue % of Wilks' Chi- sq df sig male female

clavicle mid-shaft diam (max) 0.541

Figure 5: Clavicle length, by sex Figure 6: Clavicle mid-shaft diameter (maximum),

Figure 7: Clavicle mid-shaft diameter (minimum),

Figure 8: Conoid tubercle diameter (anterior-

Figure 9: Clavicle sternal facet height (superior-

Pelvis Table 15: Sex differences in the pelvis

is not statistically significant. Dimorphism is auricular surface breadth -- 91.43%

female mean = 270.06 mm, N = 35), the

Table 16: Pelvic Model: discriminant function analysis

Eigenvalues Wilks' Lambda Standardized canonical Classification

Eigenvalue % of Wilks' Chi- sq df sig male female

Figure 10: Ilium breadth, by sex

Figure 11: Iliac crest (minimum) breadth, by sex

Figure 12: Acetabulum height, by sex

Figure 13: Pubic symphysis breadth, by sex

Figure 14: Auricular surface length, by sex

Table 18: Humeral Model: discriminant function analysis

Eigenvalues Wilks' Lambda Standardized canonical discriminant Classification

Eigenvalue % of Wilks' Chi- sq df sig male female

While the sex difference in humerus length

Table 20: Radial Model: discriminant function analysis

Eigenvalues Wilks' Lambda Standardized canonical discriminant Classification

Eigenvalue % of Wilks' Chi- sq df sig male female

Figure 23: Radius height, by sex

Figure 24: Radius mid-shaft diameter

Figure 25: Radius mid-shaft diameter (minimum), by

Table 21: Forelimb Model: discriminant function analysis

Eigenvalues Wilks' Lambda Standardized canonical discriminant Classification

Eigenvalue % of Wilks' Chi- sq df sig male female

Figure 28: Femur biomechanical neck length, by sex