AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 123:146 –155 (2004)
Effects of Different Kinds of Cranial Deformation on the
Incidence of Wormian Bones
Valerie Dean O’Loughlin*
Medical Sciences Program, Indiana University, Bloomington, Indiana 47405
KEY WORDS cultural deformation; ossicles; supernumerary bone; sutural bone;
craniosynostosis
ABSTRACT Researchers have debated whether the
presence and frequency of wormian bones (sutural bones,
supernumerary bones, and ossicles) are attributable to
genetic factors, environmental factors, or both. This re-
search examines the effects of many different kinds of
cranial deformation on the incidence of wormian bones. A
sample of 127 deformed and undeformed crania from New
World archaeological sites was examined. An undeformed
cranial sample (n ⫽ 35) was compared to the following
cranially deformed groups: 1) occipital, 2) lambdoid, 3)
annular, 4) fronto-vertico-occipital, 5) parallelo-fronto-oc-
cipital, and 6) sagittal synostosis. Three levels of degree of
cultural cranial deformation were qualitatively deter-
mined. Type and number of wormian bones along each
major suture were recorded for each cranium. Group
means were analyzed using Kruskal-Wallis one-way
Wormian bones (sutural bones, supernumerary
bones, and ossicles) are irregularly shaped bones
formed from independent ossification centers found
along cranial suture lines and fontanelles (Hauser
and De Stefano, 1989). Dorsey (1897) was among the
first to suggest that pressure on the cranium from
cultural cranial deformation may influence the inci-
dence of wormian bones. Many researchers exam-
ined this question and presented hypotheses con-
cerning wormian bone etiology. The first is that the
formation and incidence of wormian bones are pri-
marily under genetic influence, and external factors
such as cultural cranial deformation do not play a
major role in wormian bone formation (Berry and
Berry, 1967; Finkel, 1976). Berry and Berry (1967)
examined the presence (but not frequency) of
wormian bones in several skeletal populations. They
suggested that the presence of wormian bones was
genetically influenced and had minimal epigenetic
influence. Finkel (1976) even suggested that
wormian bone formation was the result of a single
gene.
A second hypothesis maintains that environmen-
tal stressors (e.g., cultural cranial deformation or
experimentally created craniosynostosis) affect the
incidence of wormian bones (e.g., Dorsey, 1897; Os-
senberg, 1970). Schultz (1923) postulated that the
formation of bregmatic bones is due to delayed clo-
© 2004 WILEY-LISS, INC.
ANOVA statistical tests to test the null hypothesis that
cranial deformation does not have an effect on wormian
bone incidence. Results indicate that all forms of cranial
deformation affect the frequency of some types of wormian
bones. In particular, all cranially deformed groups exhib-
ited significantly greater frequencies of lambdoid ossicles.
Apical, parieto-mastoid, and occipito-mastoid wormian
bones also appeared with greater frequency in some
groups of culturally deformed crania. Further, varying
degrees of cultural deformation all had more lambdoid
wormian bones than the undeformed group. These results
suggest that wormian bone development in posteriorly
placed sutures may be affected more by environmental
forces than are their anteriorly placed counterparts. Am J
Phys Anthropol 123:146 –155, 2004. © 2004 Wiley-Liss, Inc.
sure of the anterior fontanelle. Pucciarelli (1974)
stated that experimental immobilization of rat cra-
nia could positively affect the frequency of wormian
bones. Bennett (1965) examined Euro-American,
Black, and Native American crania for the presence
of lambdoid wormian bones, and hypothesized that
these bones were the result of a stressor along the
lambdoid suture. White (1996) found that fronto-
occipital modification was positively associated with
the presence of lambdoid ossicles.
Craniosynostosis is the premature fusion of one or
more of the calvarial sutures (Bixler and Ward,
1987). Craniosynostosis may be considered an envi-
ronmental stressor if, by altering skull-growth vec-
tors, it affects the presence and frequency of
wormian bones. Burrows et al. (1997) examined the
Preliminary findings were presented at the 69th Annual Meeting of
the American Association of Physical Anthropologists.
*Correspondence to: Valerie O’Loughlin, Medical Sciences Pro-
gram, Jordan Hall 104, Indiana University, Bloomington, IN 47405.
E-mail: vdean@indiana.edu
Received 9 April 2001; accepted 28 February 2003.
DOI 10.1002/ajpa.10304
Published online 10 June 2003 in Wiley InterScience (www.
interscience.wiley.com).
 CRANIAL DEFORMATION AND WORMIAN BONES
frequency of coronal and sagittal wormian bones in
rabbits which had either delayed onset or experi-
mentally created coronal suture synostosis. They
discovered that the experimentally created synos-
totic group had wormian bones that appeared after
(not prior to) the onset of cranial growth alteration,
suggesting wormian bone formation was affected by
external factors.
A third hypothesis states that while the presence
of wormian bones may be genetically determined,
external factors may influence the number of such
bones (e.g., El-Najjar and Dawson, 1977; Gottlieb,
1978; Anton et al., 1992). El-Najjar and Dawson
(1977) stated that the presence of lambdoid wormian
bones is under strong genetic control, but the num-
ber of wormian bones can be influenced by cultural
cranial deformation. Gottlieb (1978) discovered that
cultural cranial deformation had a direct effect on
increased sutural complexity in the lambdoid su-
ture, and an indirect effect on increased numbers of
lambdoid wormian bones. Anton et al. (1992) noted
that undeformed crania displayed more ossicles
than circumferentially deformed crania, but the un-
deformed group had fewer ossicles than anteropos-
teriorly deformed crania. Anton et al. (1992) hypoth-
esized that since circumferential deformation will
place compressional forces on the sutures, this com-
pression will result in fewer sutural bones. In con-
trast, anteroposterior deformation places tension
(not compression) on the lateral aspect of the cra-
nium, and the increase in tension at the sutures may
result in an increase of sutural bones.
Previous studies typically examined only a few
kinds of cultural cranial deformation (e.g., El-Najjar
and Dawson, 1977; Anton et al., 1992) or examined
effects of craniosynostosis only on sutural bone fre-
quency (Burrows et al., 1997). Other studies
grouped different types and degrees of cultural de-
formation together (El-Najjar and Dawson, 1977),
without examining the possible effects these vari-
ables could have on the incidence of wormian bones.
Some studies only examined the presence of
wormian bones along a single suture (e.g., Berry and
Berry, 1967; White, 1996), without studying the fre-
quency of these ossicles.
The present study examines the effects of many
different kinds and degrees of cranial deformation
on the incidence of different kinds of wormian bones.
The cranially deformed groups include both cultur-
ally modified crania and a craniosynostosis sample.
Comparisons are made among culturally deformed,
craniosynostotic, and undeformed groups in an at-
tempt to reach a better understanding of wormian
bone development. The research hypothesis is that
cranially deformed and craniosynostotic groups will
have different frequencies of wormian bones com-
pared to an undeformed group. In particular, it is
hypothesized that cranial deformation and cranio-
synostosis will positively influence the numbers of at
least some kinds of wormian bones. Before results
147
are presented, a brief ethnographic survey of cul-
tural cranial deformation is warranted.
ETHNOGRAPHIC INTRODUCTION TO
CULTURAL CRANIAL DEFORMATION
The term “cultural cranial deformation” is used to
describe practices that alter the shape of the cra-
nium in infancy and early childhood. Since the cra-
nium is malleable in the first years of life (Dean,
1995a), it is easy to modify head shape. Evidence of
cultural cranial deformation has existed worldwide
for several thousand years (Dingwall, 1931; Dean,
1995a). Most cultural deformational procedures be-
gin within the first few days of life, and the deform-
ing apparatus is used for approximately 6 months to
1 year. Populations in Ecuador and Peru sometimes
used the deforming apparatus for as long as 3–5
years (Dingwall, 1931; Topinard, 1879). Both male
and female children had their skull shape modified.
Cultural cranial deformation could be intention-
ally induced in a variety of ways (Dingwall, 1931;
Dean, 1995a). The vault could be tightly encircled by
bandages, producing a cylindrical or conical head
shape (e.g., in South America). Securing the skull
between two boards flattened those portions of the
head in direct contact with the boards (e.g., on the
Northwest Coast of North America). In some cul-
tures, the deformation may have been uninten-
tional, as when an infant was secured on a cradle-
board for a long period of time, consequently
flattening the back of the head (e.g., the Southwest
US).
Reasons for modifying cranium shape were varied
and sometimes culture-specific (Dingwall, 1931;
Topinard, 1879; Dean, 1995a). Northwest Coast, Co-
lumbian, and Peruvian cultures viewed males who
had deformed skulls as more brave and powerful.
Most cultures that practiced intentional molding
viewed a “molded” head as a sign of high status.
Often, slaves were not allowed to practice cultural
cranial deformation, and so their head shape was a
physical sign of their social status (e.g., Northwest
Coast). In addition, culturally modified skulls were
viewed as a sign of beauty, and the more marked the
deformation, the more beautiful the person was re-
garded as being.
It is common to see “ranges” of cranial deforma-
tion within and among cultures. Some individuals
may have had slight cranial deformation, whereas
others had more marked deformation. The length of
time the deforming apparatus was used, how tightly
the apparatus fit, and the reasons for modifying
skull shape created these “ranges” of deformation
(Dingwall, 1931; Dean, 1995a). Cranial deformation
research must not ignore these ranges, and incorpo-
rate them into the research design where possible.
MATERIALS AND METHODS
One hundred twenty-seven adult crania from the
National Museum of Natural History, the Thomas
148 V.D. O’LOUGHLIN
Gilcrease Institute of American History and Art
(Tulsa, OK), and the Indiana University collections,
representing many geographically distinct New
World archaeological populations, were analyzed. A
list of deformational groups, geographic locations
and archaeological sites of crania, and cultural/tem-
poral information is given in Table 1. Those crania
with provenience information were “historic,” or dat-
ing from no later than the 1800s. There was limited
provenience information for most crania, but several
factors (e.g., condition of remains, anecdotal infor-
mation) indicated that crania were of more recent
origin. The wide range of populations was selected
because: 1) wide sampling maximizes cranial diver-
sity and minimizes the risk of unintentionally se-
lecting for population-specific traits, and 2) no single
archaeological population examined contains a suf-
ficient number of undeformed, culturally deformed,
and craniosynostotic crania. All cranial groups (ex-
cept the lambdoid deformed group) contain speci-
mens from multiple, geographically distinct regions.
Lamboid deformed specimens came from multiple
archaeological sites but were limited to the South-
west US. Detailed ethnographic information about
the deformational practices of these regions is in
Dingwall (1931) and Dean (1995a).
Age was estimated by endocranial and ectocranial
suture closure and dental wear. This study did not
need the exact age of each cranium, but needed an
estimate as to whether the cranium belonged to a
younger (18 – 40 years) or older (40 –55⫹ years) in-
dividual. Meindl and Lovejoy (1985) noted that al-
though ectocranial suture closure scoring can have
overlap and variability, this method is still able to
provide a good general age-range estimate. As with
suture closure, dental wear evaluation was used
only as a relative measure of age, to separate
younger from older individuals. As others noted
(Hillson, 2000), dental wear varies considerably
among populations, and may not be a reliable exact
age indicator for crania from different populations.
Younger male individuals (ages 18 – 40) were se-
lected to decrease the possibility of age-related su-
ture closure and to avoid overlooking obscured
wormian bones. Ages ranged from 18 – 40 years for
all but two craniosynostotic crania (age range,
40 –55 years). Population frequencies of craniosyn-
ostosis range from 3–15 per 10,000 individuals (Co-
hen, 1986b; French et al., 1990), so control on age
and sex in the craniosynostotic group was relaxed to
maximize sample size. As a result, some female cra-
nia were included in the undeformed and occipital
deformed samples for comparative purposes. Berry
and Berry (1967) showed that sutural bones appear
in equal frequency between the sexes, so the benefit
of having a larger craniosynostotic sample size out-
weighed the risk of having a mixture of male and
female crania in three of the samples.
Type and degree of cultural cranial deformation
were qualitatively determined by visual inspection.
Deformational types included those described by
Neumann (1942) and Hrdlicka (1910): 1) occipital
deformation (n ⫽ 30), 2) lambdoid (n ⫽ 9), 3) fronto-
vertico-occipital (n ⫽ 13), 4) parallelo-fronto-occipi-
tal (n ⫽ 18), and 5) annular (n ⫽ 14) (see Fig. 1).
Occipital deformation is a vertical flattening of the
nuchal portion of the occipital bone. Lambdoid de-
formation is a flattening of the cranium around the
region of lambda. Fronto-vertico-occipital deforma-
tion is a vertical flattening of the upper portion of
the occipital, as well as an oblique flattening of the
frontal bone. Parallelo-fronto-occipital deformation
flattens the frontal region and the occipital bone
proper. The occipital bone is flattened obliquely,
whereby the frontal and occipital bones are approx-
imately parallel to each another. Annular deforma-
tion compresses the cranium cylindrically, so that
the cranium becomes ovoid.
Degree of deformation was classified as none,
slight, moderate, or marked. “No cranial deforma-
tion” was defined as no evidence of cultural cranial
deformation. “Slight deformation” was some evi-
dence of altered cranium shape, but which could
be overlooked in more casual observation. “Mod-
erate deformation” was noticeable alteration of
cranium shape. “Marked deformation” was clear
indication of drastically altered cranium shape.
Degree was determined by two or more visual
inspections by the author, using multiple compar-
ative crania from the same area. Detailed methods
of analysis for type and degree of cultural cranial
deformation were previously described (Dean,
1995a,b; O’Loughlin, 1996).
Isolated (as opposed to syndromic) craniosynosto-
ses were selected for, to minimize the possibility
that a syndrome itself caused the wormian bone
incidence, as opposed to the resulting premature
suture closure. Syndromic craniosynostosis also has
a strong genetic component, whereas isolated cra-
niosynostosis may be the result of environmental
factors, such as in utero constraint (Cohen, 1986a,
1988; Bixler and Ward, 1987). Isolated craniosynos-
toses were determined by inspecting for premature
endocranial and ectocranial sutural fusion, and not-
ing that the crania lacked certain facial anomalies
characteristic of syndromic craniosynostoses (Cohen
1986a, 1988; Bixler and Ward, 1987). All crania (n ⫽
7) displayed sagittal synostosis; two crania also had
either lambdoid synostosis or unilateral occipito-
mastoid synostosis.
An undeformed cranial sample (n ⫽ 35) from mul-
tiple New World archaeological populations was
used for comparative purposes. These populations
were different from the deformed populations, be-
cause according to the ethnographic literature (re-
viewed in Dingwall, 1931; Dean, 1995a), culturally
modifying the cranium was a practice often limited
to individuals of higher classes. Thus, undeformed
crania in culturally deformed populations were more
likely to be individuals of a different socioeconomic
setting (and perhaps a different genetic and/or nu-
 CRANIAL DEFORMATION AND WORMIAN BONES 149
TABLE 1. Geographic, site, and cultural affiliation and temporal distribution of cranial samples
Geographic location Archaeological site(s) Cultural affiliation Temporal information
Cranial group (number of individuals) Sex (if known)1 (if known) (if known)2
Undeformed (n ⫽ 35)
Montana (5) M Benton City Spokane Historic
M Tongue River Sioux Historic
M Fort Ellis Unknown Historic
M Fort Shaw Unknown Historic
M unknown Unknown Unknown
Kansas (2) M Doniphan site Unknown Unknown
M near Fort Harker Kaw Unknown
California (1) M Nicolaus Mound Unknown Unknown
Oregon (1) M Emigrant Springs Pah Ute Unknown
New Mexico (2) M Fort Wingate Navaho Late 1800s
M Fort McRae Navaho Historic
Utah (1) M Rock Grave Pah Vant Unknown
Canada (1) M Fort Good Hope Hare Historic
Arkansas (2) F Unknown Unknown Unknown
M Dickson Farm Unknown Unknown
Texas (4) M Columbia Comanche Historic
M Pecos River Comanche Historic
M Fort Concho Comanche Historic
M Fort McKarth Comanche Historic
South Dakota (5) M (3) Swan Creek site Arikara Historic
M (2) Mobridge site Possibly Arikara 1700s–1800s
Illinois (7) F Greene County Unknown Unknown
F Snyders site Woodland Woodland
M (3) Jersey County Woodland Woodland
M (2) Barry Farm Hopewell Historic
MD (1) M Ossuary 4, Ferguson Farm Mid-Atlantic seaboard Historic
Unknown location (3) F Unknown Arikara Unknown
F Unknown Chickasaw Unknown
M Unknown Unknown Unknown
Occipital deformation
(n ⫽ 30)
Peru (18) M (15) Pachacamac Pachacamac 0–1500 AD
M Chilca Unknown Unknown
M (2) Near fortress of Paramonga Unknown Middle/Late period
New Mexico (5) M (2) Guisiwa, Jemez Valley Pueblo 17th century
M Puye Tewa Unknown
M Hawikuh Zuni Unknown
M Pueblo Bonito Zuni Unknown
Arizona (4) F Unknown Unknown Unknown
M (2) McDonald’s Canyon Ancient Pueblo Unknown
M Crescent Cliff Ancient Pueblo Unknown
Arkansas (2) F Unknown Unknown Unknown
F Unknown Arikara (possibly) Unknown
Northwest Coast (1) F Shell Mound Unknown Unknown
Lambdoid deformation
(n ⫽ 9)
New Mexico (6) M Pajarito plateau Tewa Unknown
M Kwasteyukwa Pueblo Unknown
M (2) Hawikuh Pueblo Zuni Unknown
M (2) Pueblo Bonito Ruin Pueblo Unknown
Arizona (2) M Canyon del Muerto Navaho Unknown
M Near Allantown Pueblo Unknown
Utah (1) M Alkali Ridge Unknown Unknown
Fronto-vertico-occipital
(n ⫽ 13)
Washington (6) M (2) Neah Bay Makah Historic
M (3) Ft. Townsend Cowichan Historic
M Vantage Ferry Unknown Unknown
Peru (6) M Lima Unknown Unknown
M (3) Pachacamac Pachacamac 0–1500 AD
M (2) Chicama Unknown Unknown
Bolivia (1) M Cachilaya Unknown Unknown
Parallelo-fronto-occipital
(n ⫽ 18)
Washington (5) M Pacific Country Salish Historic
M (2) Puget Sound Nisqually Historic
M (2) Pacific Country, Columbia River Chinook Historic
Oregon (4) M (2) Unknown Chinook Historic
M (2) Memaloose Island Unknown Historic
Arkansas (1) M Newport Unknown Historic
Peru (4) M (2) Lima Unknown Unknown
M Candivilla Unknown Unknown
M Pachacamac Pachacamac 0–1500 AD
Dominican Republic (3) M (3) Countanza Arawak Historic
Venezuela (1) M Lake Tacarigua Unknown Unknown
(Continued)
150 V.D. O’LOUGHLIN
TABLE 1. (Continued)
Geographic location Archaeological site(s) Cultural affiliation Temporal information
Cranial group (number of individuals) Sex (if known)1 (if known) (if known)2
Annular (n ⫽ 14)
Chile (1) M Aricara Unknown Unknown
Bolivia (3) M Lake Titicaca Aymara Unknown
M Sicasica Unknown Unknown
M Cachilaya Unknown Unknown
Peru (10) M Coyungo Unknown Unknown
M (8) Santa Lucia Unknown Unknown
M Cabeza Larga cemetery Paracas Historic
Craniosynostosis (n ⫽ 7)
California (1) M Ponce Mound Unknown Unknown
Peru (3) F San Damian Unknown Historic (estimate)
M Cincos Cerros Unknown Historic (estimate)
M Cincos Cerros Unknown Historic (estimate)
Arkansas (1) M Drew County Unknown Unknown
Maryland (1) M Ossuary #4, Accokeek Middle Atlantic seaboard Historic
Alaska (1) F Nunivak Island Eskimo Historic
1
Provenience information was limited for most of these crania. Most information included just general locality, skeletal remains
recovered, and whether remains were Native American.
2
While exact dates are not known for most specimens, other information (e.g., condition of crania, history about process of obtaining
these skeletal collections) indicate that crania are not ancient, but rather are from more recent populations.

tritional setting) than culturally deformed individu- RESULTS

als.
Number and placement of wormian bones were
determined by visual inspection. Only those bones
which were clearly visible were counted. Wormian
bones were grouped according to the suture or junc-
tion of sutures where they were found (Figs. 2, 3): 1)
coronal, 2) pterionic, 3) sagittal, 4) apical (junction of
sagittal and lambdoid sutures), 5) lambdoid, 6) as-
terionic (junction of lambdoid, parieto-mastoid, and
occipio-mastoid sutures), 7) parieto-mastoid, 8) oc-
cipito-mastoid, 9) squamosal, and 10) parietal notch
(junction of the parieto-mastoid and squamosal su-
tures). (No bregmatic bones were found in this sam-
ple.) “Inca” bones were excluded from this analysis,
because their development and occurrence appear to
be the result of nonfusion of the squamous part of
the occipital to the other components of the occipital,
and are under great genetic influence (Hauser and
De Stefano, 1989).
Statistical analysis was performed using SPSS for
Windows, release 10.0.0 (SPSS, Inc., 1999). Kruskal-
Wallis one-way ANOVA tests were used to analyze
wormian bone group means, since most ossicle fre-
quency ranges were small (ranges typically were
from 0 –3). Kruskal-Wallis ANOVA is a nonparamet-
ric test that determines a “mean rank” for each
group. A higher mean rank is indicative of a higher
frequency of wormian bones in a particular group.
Probability values of P ⱕ 0.05 were considered sta-
tistically significant. Undeformed crania were com-
pared to both culturally deformed and craniosynos-
totic crania, to determine if there were differences in
the number of wormian bones among the groups.
Next, culturally deformed crania were regrouped
according to posterior degree of deformation. Unde-
formed crania then were compared to the different
degree groups to determine if the degree of deforma-
tion affected the number and placement of wormian
bones.
Table 2 lists mean number of wormian bones per
cranial group. Means were rounded to the closest
whole number. Ranges were largest for coronal
wormian bones (ranges of 0 –11 wormian bones) and
lambdoid wormian bones (ranges of 0 –22). All cra-
nially deformed groups exhibited higher mean num-
bers of lambdoid wormian bones than the unde-
formed group.
Table 3 lists Kruskal-Wallis ANOVA results com-
paring number of wormian bones among each cra-
nial group. One of the craniosynostotic specimens
(no. 276981) was eliminated from the lambdoid
wormian bone analysis because the cranium had a
lambdoid synostosis as well, reducing the craniosyn-
ostotic sample to 6. Since the craniosynostotic spec-
imens had sagittal synostosis, this group was not
used for the sagittal wormian bone analysis. Signif-
icant results were obtained for lambdoid wormian
bones and occipito-mastoid wormian bones (Table 3).
All cranially deformed groups had significantly
higher mean ranks for lambdoid wormian bones
(and thus, a higher frequency of these bones) than
the undeformed group. Two cranially deformed
groups (parallelo-fronto-occipital and annular) ex-
hibited lower mean ranks for occipito-mastoid os-
sicles than the undeformed group, and the remain-
ing groups had higher occipito-mastoid mean ranks
than the undeformed group. Culturally deformed
crania also had higher frequencies of apical bones
than the undeformed group. Table 3 also shows that
the craniosynostotic group had much higher mean
ranks for pterionic, lambdoid, occipito-mastoid, and
squamosal bones than the undeformed group. In
contrast, the undeformed group had higher frequen-
cies of coronal wormian bones.
Lambdoid and occipital deformed groups were
similar in that both had slightly greater frequencies
of coronal and apical ossicles, and much greater
 CRANIAL DEFORMATION AND WORMIAN BONES 151

Fig. 1. Cranial types used in this analysis. Crania shown are from National Museum of Natural History (Smithsonian Institution,
Washington, DC).

frequencies of lambdoid ossicles than the unde-
formed group. The parallelo-fronto-occipital and an-
nular deformed groups had greater frequencies of
coronal, pterionic, apical, squamosal, and lambdoid
ossicles than the undeformed group. In contrast, the
fronto-vertico-occipital group had lower frequencies
of coronal and pterionic wormian bones, but much
higher frequencies of almost all of the posteriorly-
placed wormian bones.
Culturally deformed crania were resorted into
simple deformed (i.e., occipital and lambdoid de-
formed) or complex deformed (i.e., fronto-vertico-oc-
cipital, parallelo-fronto-occpital, and annular)
groups for further analysis. Table 4 shows Kruskal-
152 V.D. O’LOUGHLIN
Fig. 2. Lateral view of a cranium, illustrating kinds of
wormian bones examined in this study: 1, coronal; 2, pterionic; 3,
sagittal; 4, apical; 5, lambdoid; 6, asterionic; 7, parieto-mastoid; 8,
occipito-mastoid; 9, squamosal; 10, parietal notch.
Fig. 3. Posterior view of a lambdoid deformed cranium with
multiple wormian bones: 3, sagittal wormian bone; 4, apical
wormian bone; 5, lambdoid wormian bone; 6, asterionic wormian
bone.
Wallis ANOVA results comparing the number of
wormian bones among the undeformed, simple de-
formed, and complex deformed groups. Both de-
formed groups had higher frequencies of apical and
lambdoid wormian bones than the undeformed
group. Crania in the simple deformed group had a
statistically signifcant higher mean rank of sagittal
wormian bones than the undeformed or complex
deformed groups. Since lambdoid deformed crania
(which make up part of the simple deformed group)
came from one general region, this may play a role in
these findings.
Table 5 shows Kruskal-Wallis ANOVA results
comparing posterior degree of cultural deformation
with number of wormian bones. Here, undeformed
and culturally deformed crania were regrouped ac-
cording to the degree of posterior cranial flattening
(undeformed crania were recoded as “none” for pos-
terior degree of deformation). All deformed groups
had greater frequencies of apical wormian bones,
while slight posterior flattening also was associated
with greater numbers of parieto-mastoid wormian
bones. Slight, moderate, and marked deformed cra-
nia had statistically significant higher frequencies of
lambdoid wormian bones than their undeformed
counterparts.
DISCUSSION AND CONCLUSIONS
These results indicate that all kinds of cranial
deformation (whether cultural or the result of cra-
niosynostosis) affect the frequency of certain types of
wormian bones. Wormian bone frequency may in-
crease or decrease, depending on the type and de-
gree of cranial deformation. All cranially deformed
groups had significantly greater frequencies of
lambdoid ossicles, paralleling the results of Sullivan
(1922). Apical and parieto-mastoid wormian bones
had greater frequencies in almost all deformed
groups, although these findings were not statisti-
cally significant. Note that the lambdoid, apical, and
parieto-mastoid wormian bones are in posteriorly
placed cranial sutures. Anteriorly placed wormian
bone frequencies were more similar between unde-
formed and deformed samples. In fact, two groups
(fronto-vertico-occipital and craniosynostosis) had
fewer anteriorly-placed wormian bones than the un-
deformed group.
Human wormian bone development appears to be
dependent on the location of the ossicle. I hypothe-
size that wormian bones in more posteriorly placed
sutures may be affected more by environmental fac-
tors than their anteriorly placed counterparts. Like-
wise, wormian bones in the anterior sutures appear
to be under stronger genetic control. If this hypoth-
esis is correct, it is not surprising that studies on
wormian bone incidence have yielded conflicting re-
sults.
Why would posteriorly placed wormian bones be
more influenced by environmental factors? One pos-
sibility is posteriorly placed wormian bones are
formed as a result of posteriorly placed pressure
from the cultural deforming apparatus. However,
this explanation does not explain why crania with
sagittal synostosis also have increased numbers of
posteriorly placed wormian bones. Further, Anton et
al. (1992) noted that the constriction the deforming
apparatus places on a particular suture tends to
decrease the number of sutural bones in that same
suture.
 CRANIAL DEFORMATION AND WORMIAN BONES 153
TABLE 2. Mean number of wormian bones for each cranial group1
Mean number of Wormian bones
Parietal
Cranial group Coronal Pterionic Sagittal Apical Lambdoid Asterionic Parieto-mastoid Occipito-mastoid Squamosal notch

Undeformed (n ⫽ 35) 1 (0–11) 0 (0–2) 0 (0) 0 (0–1) 3 (0–10) 1 (0–2) 0 (0–2) 0 (0–2) 0 (0–1) 0 (0–2)
Occipital (n ⫽ 30) 3 (0–10) 0 (0–2) 0 (0–4) 0 (0–1) 6 (0–16) 1 (0–2) 0 (0–2) 1 (0–5) 0 (0–3) 0 (0–2)
Lambdoid (n ⫽ 9) 2 (0–10) 0 (0–1) 0 (0–1) 1 (0–1) 7 (1–11) 0 (0–1) 1 (0–6) 1 (0–2) 0 (0–2) 0 (0–1)
Fronto-vertico-occipital 1 (0–7) 0 (0–1) 0 (0) 0 (0–1) 7 (0–22) 1 (0–2) 0 (0–1) 1 (0–3) 0 (0–1) 0 (0–1)
(n ⫽ 13)
Parallelo-fronto-occipital 2 (0–9) 0 (0–2) 0 (0) 0 (0–1) 6 (0–20) 1 (0–2) 0 (0–1) 0 (0–1) 1 (0–12) 1 (0–2)
(n ⫽ 18)
Annular (n ⫽ 14) 2 (0–6) 0 (0–1) 0 (0) 0 (0–1) 4 (0–12) 0 (0–2) 0 (0–3) 0 (0) 0 (0–2) 1 (0–2)
Craniosynostosis (n ⫽ 7) 0 (0–3) 1 (0–2) — 0 (0–1) 6 (0–14)2 1 (0–2) 0 (0–1) 1 (0–2) 1 (0–4) 1 (0–2)

1
Wormian bone means were rounded to closest whole number. Ranges for each sample are included in parentheses.
2
The only craniosynostotic specimen that did not have lambdoid ossicles was that which had lambdoid synostosis (number 276981).

TABLE 3. Kruskal-Wallis ANOVA results comparing cranial group with number of wormian bones1
Mean rank values for cranial group ANOVA values
Wormian bone Chi-square
type Undeformed Occipital Lambdoid F-V-O2 P-F-O3 Annular Craniosynostosis statistic P-value
Coronal 59.4 72.8 64.9 52.1 66.7 66.5 49.36 6.85 0.335
Pterionic 60.1 59.3 59.6 57.6 71.2 74.3 73.0 8.69 0.192
Sagittal 58.5 62.5 65.0 58.5 58.5 58.5 — 6.75 0.240
Apical 57.4 64.0 78.0 72.1 64.0 65.5 52.0 5.86 0.439
Lambdoid4 46.8 71.4 83.2 66.3 66.6 56.8 81.94 13.93 0.030*
Asterionic 61.5 71.7 41.2 69.0 73.2 50.4 58.1 10.55 0.103
Parieto-mastoid 59.9 62.2 72.3 72.0 61.4 62.8 66.6 7.45 0.281
Occipito-mastoid 60.4 65.1 77.7 82.0 54.2 48.0 74.1 16.33 0.012*
Squamosal 59.7 62.2 65.1 62.6 68.7 62.6 76.1 6.88 0.332
Parietal notch 62.7 61.1 58.1 58.5 65.2 68.3 80.4 3.75 0.710
1
Craniosynostosis group was not included in sagittal wormian bone analysis.
2
Fronto-vertico-occipital deformation.
3
Parallelo-fronto-occipital deformation.
4
Since one cranium (no. 276981) in craniosynostotic group also had lambdoid synostosis, only 6 of 7 craniosynostotic crania were used
to analyze lambdoid wormian bone frequency.
* Significant at P ⬍ 0.05.

TABLE 4. Kruskal-Wallis ANOVA results, comparing collapsed cranial groups with number of wormian bones1
Mean rank values for cranial group ANOVA values
Wormian bone Undeformed Simple deformation Complex deformation Chi-square
type (n ⫽ 35) (n ⫽ 39)2 (n ⫽ 45)3 statistic P-value
Coronal 55.3 66.3 58.2 2.82 0.245
Pterionic 57.3 56.6 65.1 3.87 0.144
Sagittal 58.5 63.1 58.5 6.26 0.044*
Apical 53.6 62.9 62.5 2.55 0.279
Lambdoid 45.5 71.5 61.3 10.67 0.005**
Asterionic 57.8 60.8 61.0 0.25 0.883
Parieto-mastoid 56.7 61.2 61.5 1.91 0.384
Occipito-mastoid 57.7 64.8 57.6 2.06 0.357
Squamosal 57.1 60.1 62.1 1.98 0.372
Parietal Notch 60.2 58.0 61.6 0.38 0.827
1
Craniosynostosis group was not included in this analysis.
2
Simple deformation group includes occipital and lambdoid deformed crania.
3
Complex deformation group includes fronto-vertico-occipital, parallelo-fronto-occipital, and annular deformed crania.
* Significant at P ⬍ 0.05.
** Significant at P ⬍ 0.01.

Another possibility relates to human brain and growth of the brain and cranium places greater
cranium development. Moss (1958) noted that the intracranial pressure on the lambdoid suture.
occipital portion of the brain grows most rapidly Greater pressure would create increased tension
after birth and positively affects the postnatal along the suture. How would this affect the cra-
growth of the posterior part of the cranium. This niosynostotic group, where the greatest compres-
finding suggests that the posterior cranium could sion is along the fused sagittal suture? Moss
be influenced more by external (epigenetic) factors (1959) stated that sagittal synostosis redirects the
than the anterior cranium. Trinkaus and LeMay growth vectors of the neurocranium in a more
(1982) suggested that this posteriorly directed anterior-posterior fashion. The end result could be
154 V.D. O’LOUGHLIN
TABLE 5. Kruskal-Wallis ANOVA results, comparing posterior degree of cultural deformation with number of wormian bones1
Mean rank values for cranial group1 ANOVA values
Wormian bone None Slight Moderate Marked Chi-square
type (n ⫽ 35) (n ⫽ 12) (n ⫽ 36) (n ⫽ 36) statistic P-value
Coronal 55.3 77.0 62.7 56.2 5.67 0.127
Pterionic 57.3 60.8 62.1 60.3 0.89 0.828
Sagittal 58.5 63.5 61.8 58.5 4.83 0.185
Apical 53.6 54.9 66.4 61.5 4.17 0.244
Lambdoid 45.5 70.2 65.4 65.3 9.05 0.029*
Asterionic 57.8 53.2 62.6 61.8 1.13 0.771
Parieto-mastoid 56.7 70.3 56.7 63.0 8.63 0.035*
Occipito-mastoid 57.7 55.9 58.7 64.9 2.01 0.570
Squamosal 57.1 55.5 63.8 60.5 4.21 0.240
Parietal notch 60.2 60.0 61.7 58.1 0.32 0.956
1
All culturally deformed groups were combined into one big group, and then resorted by degree of posterior flattening. Craniosyn-
ostosis group was not included in this analysis.
* Significant at P ⬍ 0.05.

an increased number of lambdoid wormian bones,
as seen in this study. The craniosynostotic group
had a slightly decreased frequency of coronal
wormian bones. Redirected cranial growth vectors
would not have a positive effect on coronal ossicle
frequency if these anteriorly placed ossicles are
under greater genetic control.
Per the hypothesis of Anton et al. (1992), in-
creased compression at sutures could explain why
some deformational groups had lower frequencies of
some wormian bones. For example, annular and par-
allelo-fronto-occipital deformation both place com-
pression at the occipito-mastoid sutures, and these
groups also have fewer occipito-mastoid wormian
bones. In contrast, occipital deformation does not
directly compress the entire lambdoid suture, so it
may produce tension at parts of the lambdoid suture
and thereby produce greater frequencies of lambdoid
ossicles. Lambdoid deformation compresses the
lambdoid suture directly, but since the lamboid
group was small and necessarily restricted to the
Southwest US, another unknown factor may be re-
sponsible for lambdoid ossicle formation in this
group.
Degree-of-deformation data (Table 5) show that
the posteriorly-placed lambdoid, apical, and aste-
rionic bones appear in greater frequencies in mod-
erate and marked deformed crania. With few ex-
ceptions, anteriorly placed ossicles are seen in
similar frequencies among all degree groups. Due
to sample-size limitations, the degree of deforma-
tion within a particular group (e.g., occipital)
could not be examined. As a result, type-specific
information about degree of deformation cannot be
determined. It would be interesting to know if, for
example, crania with marked occipital deforma-
tion exhibited greater frequencies of wormian
bones than crania with slight occipital deforma-
tion.
The data show that posteriorly placed wormian
bones appear in greater numbers in deformed cra-
nia. The data cannot answer whether cranial defor-
mation affects the initial presence or absence of
these ossicles. Previous animal studies yielded in-
consistent results. While Pucciarelli (1974) found
that both anterior deformation and posterior defor-
mation resulted in a similar increase of both ante-
rior and posterior sutural bones in 21 rats, 77 other
deformed rat crania exhibited no sutural bones, sug-
gesting that deformation did not affect sutural bone
presence. Burrows et al. (1997) noted in their rabbit
study that coronal wormian bones appeared after
the coronal suture was experimentally immobilized,
but as they were unable to examine the sutures in
utero, it is not clear whether wormian bones were
present before or after synostotic changes occurred.
Ultimately, the question about genetic vs. environ-
mental influence on wormian bone formation may
best be answered in the future with fetal animal
crania studies.
This study provided preliminary data and hypoth-
eses to explain wormian bone formation. More re-
search is necessary, using larger adult sample sizes,
juvenile human crania samples (Dean, 1991), and
animal studies to further test these hypotheses.
ACKNOWLEDGMENTS
Crania were provided by Indiana University
(Bloomington, IN), the Thomas Gilcrease Institute
of American History and Art (Tulsa, OK), and the
National Museum of Natural History, Smithso-
nian Institution (Washington, DC). I thank these
institutions for granting me access to the skeletal
collections and for assisting me whenever possi-
ble. I appreciate the helpful comments of the anon-
ymous reviewers of this manuscript. I thank Paul
Jamison of Indiana University for his assistance
with statistical analysis. Finally, I am indebted to
David Hunt, Physical Anthropology Collections
Manager at the Smithsonian Institution, for his
guidance and assistance with my research during
my 7-month stay.
LITERATURE CITED
Anton SC, Jaslow CR, Swartz SM. 1992. Sutural complexity in
artificially deformed human (Homo sapiens) crania. J Morphol
214:321–332.
Bennett KA. 1965. The etiology and genetics of wormian bones.
Am J Phys Anthropol 23:255–260.
 CRANIAL DEFORMATION AND WORMIAN BONES
Berry AC, Berry RJ. 1967. Epigenetic variation in the human
cranium. J Anat 101:361–379.
Bixler D, Ward R. 1987. Craniosynostosis. Handbk Clin Neurol
6:113–128.
Burrows AM, Caruso KA, Mooney MP, Smith TD, Losken HW,
Siegel MI. 1997. Sutural bone frequency in synostotic rabbit
crania. Am J Phys Anthropol 102:555–563.
Cohen MM. 1986a. History, terminology and classification of cra-
niosynostosis. In: Cohen MM, editor. Craniosynostosis: diagno-
sis, evaluation and management. New York: Raven Press, Inc.
p 1–20.
Cohen MM. 1986b. Perspectives on craniosynostosis. In: Cohen
MM, editor. Craniosynostosis: diagnosis, evaluation and man-
agement. New York: Raven Press, Inc. p 21–57.
Cohen MM. 1988. Craniosynostosis update 1987. Am J Med
Genet [Suppl] 4:99 –148.
Dean VL. 1991. The incidence of wormian bones among juvenile
crania excavated from the Schild Mississippian knolls: a case
study. Proc Indiana Acad Sci 100:113–121.
Dean VL. 1995a. Effects of cultural cranial deformation and cra-
niosynostosis on cranial venous sinus and middle meningeal
vessel pattern expression. Ph.D. dissertation, Indiana Univer-
sity, Bloomington, IN.
Dean VL. 1995b. Sinus and meningeal vessel pattern changes
induced by artificial cranial deformation. Int J Osteoarchaeol
5:1–14.
Dingwall EJ. 1931. Artificial cranial deformation: a contribution
to the study of ethnic mutilations. London: John Bale, Sons and
Danielsson, Ltd.
Dorsey GA. 1897. Wormian bones in artificially deformed
Kwakiutl crania. Am Anthropol 10:169 –173.
El-Najjar MY, Dawson GL. 1977. The effect of artificial cranial
deformation on the incidence of wormian bones in the lambdoi-
dal suture. Am J Phys Anthropol 46:155–160.
Finkel DJ. 1976. Wormian bone formation in the skeletal popu-
lation from Lachish. J Hum Evol 5:291–295.
French LR, Jackson IT, Melton LJ. 1990. A population-based
study of craniosynostosis. J Clin Epidemiol 43:69 –73.
Gottlieb K. 1978. Artificial cranial deformation and increased
complexity of the lambdoidal suture. Am J Phys Anthropol
48:213–214.
Hauser G, De Stefano GF. 1989. Epigenetic variants of the human
cranium. Stuttgart: E. Schweizerbart Verlagsbuchhandlung.
155
Hillson S. 2000. Dental pathology. In: Katzenburg MA, Saunders
SR, editors. Biological anthropology of the human skeleton.
New York: Wiley-Liss, Inc. p 249 –286.
Hrdlicka A. 1910. Artificial deformations of the human cranium,
with especial reference to America. Int Cong Americanists Ac-
tas 17:147–149.
Meindl RS, Lovejoy CO. 1985. Ectocranial suture closure: a re-
vised method for the determination of skeletal age at death
based on the lateral-anterior sutures. Am J Phys Anthropol
68:57– 66.
Moss ML. 1958. The pathogenesis of artificial cranial deforma-
tion. Am J Phys Anthropol 16:269 –286.
Moss ML.1959. The pathogenesis of premature cranial synostosis
in man. Acta Anat (Basel) 37:351–370.
Neumann GK. 1942. Types of artificial cranial deformation in the
eastern United States. Am Antiq 3:306 –318.
O’Loughlin VD. 1996. Comparative endocranial vascular changes
due to craniosynostosis and artificial cranial deformation. Am J
Phys Anthropol 101:369 –385.
O’Loughlin VD. 2000. Do different kinds of cranial deformation
affect the incidence of wormian bones in human crania? Am J
Phys Anthropol [Suppl] 20:134 –135 [abstract].
Ossenberg NS. 1970. The influence of artificial cranial deforma-
tion on discontinuous morphological traits. Am J Phys An-
thropol 33:357–372.
Pucciarelli HM. 1974. The influence of experimental deformation
on neurocranial wormian bones in rats. Am J Phys Anthropol
41:29 –38.
Schultz AH. 1923. Bregmatic fontanelle bones in mammals. J
Mamm 4:65–77.
SPSS, Inc. 1999. SPSS base 10.0 syntax reference guide. Chicago:
SPSS, Inc.
Sullivan LR. 1922. The frequency and distribution of some ana-
tomical variations in American crania. Anthropol Pap Am Mus
Nat Hist 23:203–258.
Topinard P. 1879. Des deformations ethniques du crâne. Rev
Anthropol 2:496 –506.
Trinkaus E, LeMay M. 1982. Occipital bunning among later
Pleistocene hominids. Am J Phys Anthropol 57:27–35.
White CD. 1996. Sutural effects of fronto-occipital cranial modi-
fication. Am J Phys Anthropol 100:397– 410.