AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 00:00–00 (2014)
Comparability of Multiple Data Types From the Bering
Strait Region: Cranial and Dental Metrics and
Nonmetrics, mtDNA, and Y-Chromosome DNA
Brianne Herrera,1* Tsunehiko Hanihara,2 and Kanya Godde3,4
1
Department of Anthropology, The Ohio State University, Columbus, OH
2
Department of Anatomy and Biological Anthropology, Saga Medical School, Saga, Japan
3
Sociology and Anthropology Department, University of La Verne, La Verne, CA
4
Department of Anthropology, University of Tennessee, Knoxville, TN
KEY WORDS mantel tests; biological distance; aleut; inuit; procrustes; kinship matrix
ABSTRACT Different data types have previously
been shown to have the same microevolutionary pat-
terns in worldwide data sets. However, peopling of the
New World studies have shown a difference in migration
paths and timings using multiple types of data, spurring
research to understand why this is the case. This study
was designed to test the degree of similarity in evolu-
tionary patterns by using cranial and dental metric and
nonmetric data, along with Y-chromosome DNA and
mtDNA. The populations used included Inuits from
Alaska, Canada, Siberia, Greenland, and the Aleutian
Islands. For comparability, the populations used for the
cranial and molecular data were from similar geographic
regions or had a shared population history. Distance, R
and kinship matrices were generated for use in running
Mantel tests, PROTEST analyses, and Procrustes analy-
Combining different types of data for statistical analy-
sis is useful for inferring patterns of population dynam-
ics and history. Genetic drift, gene flow, and natural
selection are well-known biological and evolutionary
influencers of cranial shape and form that are passed
down through generations (Relethford and Blangero,
1990; Relethford, 2001; Stojanowski, 2004). Because of
this, comparing osteological and molecular data has had
interesting results in investigations ranging from kin-
ship organization during the Late Stone Age (Haak
et al., 2008) to finding familial groups within ancient
Mongolians (Ricaut et al., 2010). Gonz alez-Jose et al.
(2008) have also been successful in combining geometric-
morphometrics and genetic data for testing different
models for the human migration into the Americas. Sim-
ilarly, Gonz
alez-Jose et al. (2004) used cranial measure-
ments and Alu-insertions to find correlations with
genetic and phenotypic distances.
The analysis performed here is based on a similar
idea. We used osteological metric and nonmetric data,
dental metric and nonmetric data, as well as mitochon-
drial DNA (mtDNA) haplogroup frequencies and Y-
chromosome allele frequency data from different contem-
porary, indigenous Arctic and Aleutian populations in
order to compare and detect population patterns. Specifi-
cally, we are interested in combining multiple biological
data types in hopes that the methodology will aid in bio-
archaeological investigations, in particular, peopling of
the New World studies. Because this work will not
address temporal or spatial components associated with
Ó 2014 WILEY PERIODICALS, INC.
ses. A clear patterning was seen, with the craniometric
data being most highly correlated to the mtDNA data
and the cranial nonmetric data being most highly corre-
lated with the Y-chromosome data, while the phenotypic
data were also linked. This patterning is suggestive of a
possible male or female inheritance, or the correlated
data types are affected by the same or similar evolution-
ary forces. The results of this study indicate cranial
traits have some degree of heritability. Moreover, com-
bining data types leads to a richer knowledge of biologi-
cal affinity. This understanding is important for
bioarchaeological contexts, in particular, peopling of the
New World studies where focusing on reconciling the
results from comparing multiple data types is necessary
to move forward. Am J Phys Anthropol 000:000–000,
2014. VC 2014 Wiley Periodicals, Inc.
these populations and data types, it is not appropriate to
directly examine these results for human migration to
the New World. Instead, we hope this information will
provide a tool for investigators wishing to analyze multi-
ple data types in relation to hypothesis testing for peo-
pling of the New World. Because of the large variety of
data types used for migration studies, it is important to
understand the relationship of each data type to each
other for researchers to reconcile the results.
BACKGROUND
Greenland
The first migration into Greenland is thought to have
occurred via the Canadian High Arctic islands (Mel-
gaard, 1976) around 4500–4000 BP (Helgason et al.,
2005). Once in Greenland, the original migrants split
Additional Supporting Information may be found in the online ver-
sion of this article.
*Correspondence to: Brianne Herrera, 825 N 4th St #208, Colum-
bus, Ohio 43215, USA. E-mail: brianne215@gmail.com
Received 16 August 2013; revised 3 March 2014; accepted 4
March 2014
DOI 10.1002/ajpa.22513
Published online 00 Month 2014 in Wiley Online Library
(wileyonlinelibrary.com).
2 B. HERRERA ET AL.
and went down the east and west coast until making it
to the southern portion. This splitting led to the develop-
ment of different cultural groups: Independence I—Saq-
qaq, Pre-Dorset, Independence II—Dorset, and Thule
(Rasmussen et al., 2010). The Thule later gave rise to
the modern day Alaskan Yupik and Inupiat, as well as
Canadian and Greenlandic Inuit (Gilbert et al., 2008).
Ancient DNA analysis of a 4,000-year-old Greenlander
from Qeqertasussuk, Greenland shows evidence that he
was genetically distinct from the ancestors of modern
day Native Americans and Inuit, but shares ancestry
with Arctic north-east Asians (Rasmussen et al., 2010).
There may have been a few conflicts with the Native
Greenlanders before the Norse abandoned their Green-
land dwellings due to lack of regular contact with
Europe, lack of resources, declining population, unstable
climate, and overgrazing (Berglund, 1986). There was a
later migration of the Danish, and historical evidence
indicates that the Danish migration consisted of mostly
men who are known to have married native Green-
landers (Gad, 1971).
Canada
Paleo-American sites in Canada indicate the first peo-
ple arrived at 4500 BP, and over the immediately subse-
quent 500 years, population levels were rising (Savelle
and Dyke, 2002). There was a peak in the population
between 4000 and 3800 BP, followed by a sharp decline
within a few decades of 3800 BP (Savelle and Dyke,
2002). Large populations existed until 3200 BP and then
declined further, followed by a resurgence during 3000–
2800 BP (Savelle and Dyke, 2002). Afterward, another
decline occurred, possibly bringing the population down
to zero around 2800–2600 BP. A later culture, the Dor-
set, developed in the Canadian Arctic around 500 BC
(2500 BP) by the descendants of the people who had
arrived there from Alaska between 1,500 and 1,000
years earlier (Savelle and Dyke, 2002).
Siberia
The ancestral Native American population is thought
to have originated somewhere near the Lake Baikal
region and the Altai Mountains (Malyarchuk et al.,
2011; Battaglia et al., 2013). These groups are tradition-
ally Turkic-speaking tribes (Zlojutro et al., 2009b). After
Russian contact, the Yakut expanded beyond the Yaku-
tian valley and into territory occupied by the Yakaghirs,
Evenks, and Evens (Zlojutro et al., 2009b). Beginning
around early 20th century, the Siberian populations
were reduced to only the southern and northeastern por-
tion of the Chukchi Peninsula (Starikovskaya et al.,
1998). This reduction in the Siberian population was
likely due to disease from European explorers, with
many of the surviving Chukchi and Yupik people forcibly
relocated (Holzlehner, 2011). Since this occurrence, the
Siberian region has been inhabited by three different
Yupik-speaking tribes, each of which occasionally inter-
bred with various indigenous groups from neighboring
islands, such as Little Diomede Island or St. Lawrence
Island (Starikovskaya et al., 1998).
Alaska and Aleutian Islands
The Thule Tradition in Alaska is thought to have
started in Northern Alaska, eventually spreading to the
Alaskan Peninsula sometime around 1000 BP. This
American Journal of Physical Anthropology
migration of Thule replaced the Norton Culture, who
were present along the Bering Sea Coasts, starting
around 2500 years BP (Britton et al., 2013). During the
late Siberian Paleolithic stage, a group of people crossed
into the Americas from Asia via the Bering land bridge.
By 8700 BP, they eventually made it to the Aleutian
Islands, a group of islands extending westward from the
Alaska Peninsula (Smith et al., 2009). In 1791, the Rus-
sians made contact with the Aleuts, and after a few con-
flicts with the native people, the Russians and Aleutians
lived in relative harmony. After the US purchased
Alaska from Russia in 1867, the Aleutians underwent a
violent series of contact (e.g., invasions, internment
camps during WWII) from the United States and Japan
(Sepez et al., 2007; West et al., 2010). The sale also
brought Scandinavian and Western European fur trad-
ers and fisherman (Zlojutro et al., 2009a; Graf et al.,
2010; Rey et al., 2010). Intermarriage between Euro-
peans and the native Aleut population were high, par-
tially due to a law imposed by the US that required a
man to be married to an Aleutian to hunt animals typi-
cally used for fur (Reedy-Maschner, 2010). Further, the
Russian governor encouraged males to marry Aleutian
women as a way to increase fertility (Rubicz et al.,
2010b). The resulting gene flow has been detectable in
genetic studies (Rubicz et al., 2003, 2010b; Zlojutro
et al., 2006; Crawford et al., 2010).
Archaeologically, the Aleutian Islands have been a
great source of interesting and useful finds, such as
numerous mummies with soft tissue and clothing still
present (Candela, 1939; Zimmerman and Aufderheide,
1984; Veltre and Smith, 2010). The earliest site dates to
9,000 years ago, but there are only a few sites dating
around this time frame (Veltre and Smith, 2010). How-
ever, archaeologists have found about 1,000 sites dating
from 3,000 to 4,000 years ago. These sites tend to be
well preserved, allowing archaeologists to learn about
the Aleut lifestyle. Hrdlička collected skeletal material
from the Aleutian Islands during one of his expeditions.
In 1945, he published a paper suggesting an original
migration founding the Aleutian isles, and then a later
migration from the mainland of Alaska replacing the
original founding population (Hrdlička, 1945). This
theory was based on the presence of two distinct cranial
morphologies seen in his collection, but, unfortunately,
his excavation did not include stratigraphy or specific
geographic location of the skeletons (Smith et al., 2010).
Genetic studies have found evidence both for and
against this theory (Raff et al., 2010; Rubicz et al.,
2010a; Smith et al., 2010) with no consensus as of yet.
Integration of multiple data types
We expect our data types to demonstrate similarities
when considering among and between group affinities.
This line of thought is based on previous work by Ricaut
et al. (2010) and Gonz alez-Jose et al. (2008). Ricaut
et al. (2010) conclude that nonmetric traits are not as
useful for kinship patterns as genetics, but are able to
act as a proxy for genetic markers when studying among
or between group affinities. Even though our work will
be utilizing more data types (cranial metric and nonmet-
ric, dental metric and nonmetric, Y-chromosome DNA,
and mtDNA) than the Ricaut et al. (2010) article, we are
still expecting the same results in terms of group affin-
ities. Gonzalez-Jose et al. (2008) propose that models for
the human dispersion into the Americas should be based
 COMPARABILITY OF MULTIPLE DATA TYPES
on both morphology and genetic data. Furthermore, this
project is justified in light of Hubbe et al. (2009) who
state that cranial morphology can be used to infer phylo-
genetic patterns and information.
On the basis of the description of neutral theory by
Roseman and Weaver (2007), where the genetic and phe-
notypic distances are expected to be similar, we expect
to find similarities between the principal components/
coordinates of our genetic and cranial data. Harvati and
Weaver (2006) similarly suggest that cranial morphology
can retain a population history signal, allowing
researchers to track neutral genetics. However, a poten-
tial complication arises when facial shape is considered
because the face appears to retain climatic features
(Hanihara 1993, 1996). This climatic signal may be pres-
ent in arctic populations, and if so, could alter our
results. Counter to this conclusion by Harvati and
Weaver (2006), Hanihara (1997, 2000) and Perez et al.
(2007) find that facial morphology can indeed illuminate
population structure and history.
MATERIALS AND METHODS
The data sets in this project were comprised of cranio-
metric, cranial nonmetric, dental metric, dental nonmet-
ric, Y-chromosome, and mtDNA. A map of the locations
for each population from which each data type was
obtained is provided in Figure 1. The map was made
with data from Natural Earth and constructed in Cran
R (R Development Core Team, 2005), using the packages
maptools (Bivand et al., 2013), maps (Becker et al.,
2013), and mapproj (McIlroy et al., 2013). To test our
null hypothesis that all data types provide an equivalent
Fig. 1. Map of world regions represented by the various
populations utilized in this study.
TABLE 1. Howells and Martin cranial measurements used. Measurements starting with an “M” denote that Martin’s guidelines
were followed instead of Howells
Measurement
abbr. Measurement description
a,b
GOL Maximum cranial length
NOLa,b Nasion-opisthocranion
BNLa,b Cranial base length
XCBa,b Maximum cranial breadth
M9a Minimum frontal breadth
AUBa,b Biauricular breadth
ASBa,b Biasterionic breadth
BBHa,b Basion-bregma height
M26a Sagittal frontal arc
M27a Sagittal parietal arc
M28a Sagittal occipital arc
FRCa,b Nasion-bregma chord
PACa,b Bregma-lambda chord
a
Represents measurements used in matched data sets.
b
Represents measurements used in unmatched sets.
3
estimation of biological affinity, kinship matrices were
generated from each separate data type. We followed the
protocol set forth by Spradley (2006) who compared kin-
ship matrices among craniometric and allele data using
RMET 5.0 (Relethford, 2003) and Kship (Jantz, n.d.).
Cranial and dental data
Data from cranial and dental markers in contempo-
rary samples were collected by the second author (TH).
This data set is rich in comparative qualities as many
crania were observed for each of the four phenotypic
data types: craniometric, cranial nonmetric, dental met-
ric, and dental nonmetric, with the craniometric data
represented by Howells’ and Martin’s measurements
(Table 1) (Martin and Saller, 1957; Howells, 1973). Thus,
the nature of the datasets allowed us to create smaller
subsets comprised of identical individuals who had been
observed/measured for each of the data types.
Metric data and nonmetric data are subject to differ-
ent variable selection methods to contend with the
attributes of each data type. However, as the goal of this
article is comparability of the different data types, simi-
larity in variable selection techniques was also necessi-
tated. Measurements were scarce in several of the
samples included in this analysis. RMET 5.0 (Relethford,
2003) does not process missing data, and thus either
estimation of missing data must be undertaken, or the
missing data eliminated. Traditionally, missing data is
estimated. However, as the purpose of this article is
methodological and to provide correlation estimates from
comparable raw data sources, estimation of missing data
could potentially bias the results and was avoided. Thus,
variables and cases with excessive missing data were
eliminated.
Cranial and dental nonmetric data are also subject to
a different set of data selection procedures. First, bilat-
eral traits must be counted in an appropriate manner.
We followed the procedure set forth in Konigsberg (1987)
where both sides were observed and asymmetric traits
were randomly selected as to which side was utilized in
statistical analyses. Variables with excessive amounts of
missing data were eliminated, as were individual cases
presenting an excessive pattern of missing values, in
order to approximate a similar data set to the metric
Measurement
abbr. (cont’d) Measurement description
a,b
OCC Lambda-opisthion chord
M43a Breadth between
frontomalare temporale
DKBa Interorbital breadth
M51a Orbital breadth
OBHa Orbital height
NLBa Nasal breadth
NLHa Nasal height
MDBa Mastoid width
M431a Frontal chord
M43Ca Frontal subtense
XFBb Maximum frontal breadth
BPLb Basion prosthion length
NPHb Nasion prosthion height
American Journal of Physical Anthropology
4 B. HERRERA ET AL.

data (these are the unmatched data sets). Once this pro- are located in Table 2. The total sample size for the cra-
cedure was followed, the metric and nonmetric data sets nial metric data is 708 individuals and the sample size
were compared and reduced further to create data sets for the cranial nonmetric data is 766 individuals, as
with identical individuals across populations, samples, shown in Table 3. The sample size for the dental metric
and data types (deemed the matching data sets). Thus, and nonmetric data is 202 and 210 individuals, respec-
there are two types of data we worked with: 1) unmatch- tively. This is shown in Tables 4 and 5.
ing data sets where the individuals included differed
from data type to data type, and 2) matching data sets Y-chromosome data
where data types were extracted from the exact same
individuals. The final list of the Howells’ and Martin’s The Y-chromosome contains numerous genes, is only
cranial measurements utilized are listed in Table 1, and paternally inherited, and is only carried by (or present in)
the variables retained for the cranial nonmetric datasets males. Data for the Y-chromosome can be found in many
forms: short tandem repeats (STRs), single-nucleotide
polymorphisms (SNPs), haplotypes, haplogroups, and
TABLE 2. The cranial nonmetric traits utilized in the matched others (Karafet et al., 1999; Ruiz-Linares et al., 1999;
and unmatched analyses Wozniak et al., 2007). Our particular study utilizes allele
frequencies from Y-linked STRs. All the data used were
Measurement from previously published data, shown in Table 6.
Measurement description abbr.
Also, as numerated in Table 6, a total of 913 individuals
Nasofrontal and frontomaxillary sutures NFSa were used in this analysis. The frequencies were calcu-
Medial palatine canal MPCa lated by hand in Microsoft Excel (Microsoft, 2012),
Hypoglossal canal bridging HGCBa,b which were then put into the program Kship (Jantz,
Condylar canal CCAa,b n.d.). Using this program, kinship and genetic distance
Tympanic dehiscence TDa,b matrices were found.
Precondylar tubercle PCTa
Ovale-spinosum confluence OSCa
Supraorbital foramen SOFa,b Mitochondrial DNA
Transverse zygomatic suture vestige TZSb
Ossicle at lambda OLa
Mitochondrial DNA (mtDNA) is only maternally inher-
Interparietals (os incae) IPa ited, but is present in both males and females. It is com-
Biasterionic suture BASa,b monly used in genetic studies because of its fast
Accessory mental foramen AMFa mutation rate. This allows researchers to more easily
Occipitomastoid bone OMBa track new heritable mutations, which are subsequently
passed along to offspring. These minute differences allow
a
Represents traits used in matched analyses. researchers to more easily distinguish between groups of
b
Represents traits used in unmatched analyses. people (Schurr, 2002, 2004). The data used for the

TABLE 3. Cranial metric and nonmetric data, showing source population, number of individuals location of samples, and the
sources
Final Na Final Na
Population Original Na unmatching matching Location Source
Alaska1 135 and 146 114 and 146 112 and 112 Alaskan Yupik Eskimos from: Togiak; TH
Mumtrak; Tanunak; Bethel; Hooper Bay;
Iliamna Lake; Yukon; Kwichak; Nushagak
and Kuskokwim River Basin; St Lawrence
Island
Alaska2 107 and 107 101 and 107 101 and 101 Alaskan Inupiat Eskimos from: Seward TH
Peninsula, Shishmaref, Wales, Postolik,
Nome, Little Diomede Island, Point Hope
Alaska3 81 and 125 54 and 125 54 and 54 Alaska Eskimos from Point Barrow, Utkia- TH
vik, Nixerak.
Siberia 23 and 21 22 and 21 20 and 20 Port Providence, Plover Bay, Puoten, Indian TH
Point, NE Siberia.
Canadian 30 and 28 25 and 28 25 and 25 Pond’s Inlet, Sculpin Island, Labrador, TH
Manico Point, Southampton Island
Cumberland Golf, Buffin Island
Greenland 112 and 115 96 and 112 93 and 93 Smith Sound, Port Foulke, Western District, TH
Nuussuaq; Ikertok Fiord, North Star Bay,
Saunders Island
Aleut1 102 and 102 61 and 102 60 and 60 Amaknak, Unalaska, Unga, Umnak, Wislow, TH
and Shiplock Islands/Fox Islands of Aleu-
tian Island chain.
Aleut2 71 and 60 26 and 60 26 and 26 Samalga Island Quad, Kagamil Island. TH
Aleut3 13 and 13 10 and 12 10 and 10 Kanaga, Amlia, Adak, and Atka Islands; TH
Andreanof Islands.
Aleut4 18 and 29 14 and 18 13 and 13 Amchitka and Kiska Islands; Rat Islands. TH
Aleut6 16 and 20 8 and 20 8 and 8 Aleutian Island Chain—unknown locality TH

a
The first number represents the sample size from the craniometric data, the second number represents the sample size from the
cranial nonmetric data.

American Journal of Physical Anthropology

 COMPARABILITY OF MULTIPLE DATA TYPES 5
mtDNA portion of the analysis were obtained from pre- ple size and pedigree information can bypass the poten-
viously published data, shown in Table 7. A total of tial bias introduced from phenotypic data.
1,630 individuals were used for this portion of the analy- The heritability of cranial nonmetric traits has been
sis. The data used included information on the frequen- thoroughly investigated in the animal and human litera-
cies of haplogroups A, B, C, D, and “other.” ture with estimates of heritability approximately propor-
tional to craniometrics (Gr€ uneberg, 1952; Cheverud,
1981, 1988; Cheverud and Buikstra, 1981a,b, 1982;
Statistical analysis McGrath et al., 1984; Sjïvold, 1984; Richtsmeier and
McGrath, 1986). In 2006(b), Carson calculated cranial
Heritability of phenotypic data is an important consid-
nonmetric heritability with maximum likelihood estima-
eration when looking at comparability of data types.
tion, a new statistic to be applied to the problem. Her
Heritabilities differ from population to population and results suggested that heritabilities in cranial nonmetrics
environment to environment. Cranial metrics have were much lower than previously thought. Despite these
enjoyed a long established heritability value, 0.55 findings, Ricaut et al. (2010) still found concordance
(Devor, 1987), utilized in many craniometric investiga- between molecular and cranial nonmetric data, indicating
tions, across different populations. More recently, Carson heritability was strong, at least in the Mongolian popula-
(2006a) and Martınez-Abadıas et al. (2009) reexamined tion they investigated. To combat the potential issues
heritability in craniometrics, finding lower and compet- with heritability here, large sample sizes were selected
ing heritability values in different regions of the skull. and we invoked a narrow heritability of 1 for phenotypic
Moreover, issues with concordance of P (phenotypic) and data, as this is considered the conservative approach (Rel-
G (genetic) covariance matrices were noted, indicating ethford and Blangero, 1990; Relethford, 1994).
that phenotypic data are not a perfect representation of The craniometric and dental metric data were run
genetic data (Martınez-Abadıas et al., 2009). However, through RMET 5.0 (Relethford, 2003) to obtain an R
Martınez-Abadıas et al. (2009) conclude sufficient sam- matrix, or the variance covariance matrix of the data.
An R matrix further represents an estimate of genetic
TABLE 4. Dental metric data, showing source population, number kinship that details the pattern of affinity among groups
of individuals, location of samples, and the sources (Harpending and Jenkins, 1973). Relethford and Blan-
gero (1990) and Relethford et al. (1997) extended this
Population N Location Source
coefficient of kinship for multivariate application of
Alaska 66 Nunivak Island; Bering Sea TH quantitative traits. This R matrix is derived from pheno-
Greenland 58 Smith Sound, Port Foulke; TH typic data. For populations i and j, the elements of the
Disco Island; Upernatic; matrix are represented by the following equation:
Wolstenholme Sound
Aleutian 78 Unalaska, Unga, Umnak, TH
Shiplock, Amaknak, Kagamil, ðpi 2p Þðpj 2
pÞ
Kanaga, Amlia, Adak, Atka, rij 5
 ð12
p pÞ
Amchitka, Kiska, Attu, Agatu
Islands
where
X
TABLE 5. Dental nonmetric data, showing source population, p
5 wi pik
number of individuals, location of samples, and the sources
Population N Location Source with wi representing the ratio of census size of popula-
tion i to the total census size of all groups, pik is the fre-
Alaska 80 Alaska Eskimos from Point TH quency of one allele at locus k, and the summation is
Barrow, Seward Peninsula, over all groups.
Kialegak, St Lawrence Island, R matrices were also obtained from the cranial and
and S Alaska
Greenland 54 Ikertok Fiord, Smith Sound, Port TH
dental nonmetric data. These R matrices utilize a codiver-
Foulke, Mugsuak Peninsula, gence matrix derived from the biological distance of
Disco Island, Wolstenbolme Mahalanobis D2 with a tetrachoric matrix (Konigsberg,
Sound, Upernavic, Sepulchre, 2006), which provides a distance matrix for discrete data.
Issungvak The equation for Mahalanobis D2 with a tetrachoric
Aleutian 76 Aleutian Island Chain TH matrix (Williams-Blangero and Blangero, 1991) is:

TABLE 6. Y-chromosome data, showing source population, number of individuals, location of samples, and the sources
Population N Location Source
Aleutian (Aleut 1) 137 Unalaska, St. George, St. Paul, Bering Island, Akutan, False Zlojutro, 2006
Pass, King Cove, Nelson Lagoon, Sand Point
Alaskan—Yakut 150 W/SW region of Alaska Davis et al., 2011
(Alaska 1)
Alaskan—Inupiat 151 NW coast of Alaska, North Slope Borough region Davis et al., 2011
(Alaska 2)
Greenland—Inuit 70 Nanortalik, Nuuk, Ilulissat, Uummannaq, Upernavik, and Bosch et al., 2003
Ittoqqortoormiit
Greenland 272 Aasiaat, Llulissat, Kangaatsiaq, Maniitsoq, Nanortalik, Nar- Hallenberg
saq, Nuuk, Paamiut, Qaanaaq, Qaqortoq, Qasigiannguit, et al., 2009
Sisimiut, Tasiilaq, Upernavik, and Uummannaq

American Journal of Physical Anthropology

6 B. HERRERA ET AL.
TABLE 7. MtDNA data showing source population, number of individuals, location of samples, and the sources
Population N Location Source
Aleutians (Aleut 1) 198 Atka, Umnak Island, Unalaska, St. George and St. Rubicz et al., 2003
Paul Islands, Anchorage
Aleutians (Aleut 2) 78 Pribilof Island—St. Paul Merriwether et al., 1995
Aleutians (Aleut 3) 35 Bering Island of the Commanders Zlojutro et al., 2006
Greenland Eskimo 98 Nanortalik, Nuuk, Ilulissat, Uummannaq, and Saillard et al., 2000;
Upernavik, Ammassalik and Ittoqqortoormiit Shields et al., 1993
Siberian Eskimo 90 Chukotka, Nganasan of the Taimir Peninsula, Lower Derbeneva et al., 2002;
Amur River in Siberia, Chukchi Peninsula. Chukchi Shields et al., 1993;
and Siberian Eskimos collected in New Chaplino, Starikovskaya et al., 1998
Sireniki, Provideniya, and Anadyr.
Pre AD 1000 Aleuts (Aleut 4) 8 Umnak Island Smith et al., 2009
Post AD 1000 Aleuts (Aleut 5) 12 Umnak Island Smith et al., 2009
Living Aleuts (Aleut 6) 58 Umnak Island Smith et al., 2009
Canada 96 Cambridge Bay (Victoria Island)—Kitikmeot region of Volodko et al., 2008
Nunavut in Canada
Greenland 385 Ammassalik, Aasiaat, Ilulissat, Maniitsoq, Nanortalik, Volodko et al., 2008
Narsaq, Nuuk, Paamiut, Qaqortoq, Qasiqiannguit,
Qeqertarsuaq, Sisimiut, Uummannaq, and Qaanaaq
Alaska1 (Yupik) 25 Akutan, False Pass, King Cove, Nelson Lagoon, and Zlojutro et al., 2009b (from
Sand Point JH Simonson and GH
Shields, unpublished data)
Alaska2 (Inupiat) 255 Kodiak Island, St. Lawrence Island, and SW Alaska Merriwether et al., 1995
Asian (Siberia) 126 Naukan—collected in Lavrentiya (Chukotskiy district, Volodko et al., 2008
Chukotkan autonomous Region)

0
d2ij 5ðzik 2zjk Þ T 21 ðzik 2zjk Þ
where zik is the threshold for the frequency of trait k in
group i, zjk is the threshold for the frequency of trait k
in group j, and T is the tetrachoric matrix. A narrow
heritability of 1 was assumed to be consistent across
metric and nonmetric analyses. These distance matrices
(cranial nonmetric and dental nonmetric) were computed
in Fortran95 with programming provided by Dr. Lyle
Konigsberg (personal communication). The C and R mat-
rices were generated in R (R Development Core Team,
2005) with scripts provided by Dr. Lyle Konigsberg (per-
sonal communication). A detailed explanation of this
methodology is found in Konigsberg (2006).
All R and kinship matrices were subject to ordination,
via a principal coordinates analysis (PCO). The eigenvec-
tors were scaled by the square root of their individual
eigenvalues. Each kinship or R matrix was subject to a
series of two-way Mantel tests (Mantel, 1967) comparing
it to another kinship or R matrix computed from a differ-
ent data type. The two-way Mantel tests allow for the esti-
mation of the correlation between two matrices and
provide a P value representing the significance of the cor-
relation. The Mantel tests were completed in R using the
vegan package (Oksanen, 2013; Oksanen et al., 2013).
There were ten combinations of data (the matching and
unmatching sets of 1) cranial metric and nonmetric, 2)
cranial metric and Y-chromosome DNA, 3) cranial metric
and mtDNA, 4) cranial nonmetric and Y-chromosome
DNA, 5) cranial nonmetric and mtDNA) used for the Man-
tel tests, and subsequent statistical analyses.
The various data types have different units, which
limits a direct comparison. To adjust for this, a Pro-
crustes analyses was performed in R using the vegan
package (Oksanen, 2013; Oksanen et al., 2013). A Pro-
crustes analysis is a shape analysis that does a linear
transformation, which can include scaling, reflection,
rotation, or translation of the elements in a matrix to
best fit the orientation of a second matrix. Additionally,
it minimizes the sum of squares deviations, or the m2
term. In this case, the Procrustes analysis was per-
formed on the principal coordinates for different combi-
nations of two sets of data, as in Konigsberg and
Herrmann (2001). Like Konigsberg and Herrmann
(2001) and Konigsberg and Ousley (1995), the resultant
transformed principal coordinates were subsequently
graphed using scatterplot3D in R (Ligges and M€ achler,
2003). R scripts written for this manuscript will be made
available for download at: https://sites.google.com/site/
drgodde/r-scripts.
To determine whether m2 is smaller than what would
occur by chance, a PROTEST analysis was run using the
vegan package in R (Oksanen, 2013; Oksanen et al.,
2013). PROTEST has enjoyed application in ecology/evo-
lution studies (e.g., D’Anatro and Lessa, 2011), and more
recently in anthropology (Relethford, 2009), making it
an established tool in population genetics. The PRO-
TEST performs permutations on the data sets and deter-
mines concordance between the data sets more
powerfully than the Mantel test (Peres-Neto and Jack-
son, 2001). In other words, if the Mantel test shows no
statistical significance in concordance between data sets
but the PROTEST does, then there is an underlying
relationship between the two sets that was undetectable
by the Mantel test.
The Procrustes analysis can be informative about the
variation of the shape of the data set, but not the indi-
vidual points (in this case populations) and should not
be used for such purposes. The Procrustes analysis also
minimizes the shape differences among the different
data types and uses a least-squares analysis to distrib-
ute the variance equally among all of the points. If vari-
ation is needed for the individual points rather than the
shape of the whole set, a Euclidean Distance Matrix
Analysis (EDMA) should be used instead (Lele, 1993).
RESULTS
The dental data was eliminated from the analysis due
to small sample size and the subsequent failure of the

American Journal of Physical Anthropology

 COMPARABILITY OF MULTIPLE DATA TYPES 7
TABLE 8. All correlation and P values from the Mantel tests and PROTESTs performed
Mantel Mantel two-tailed PROTEST PROTEST
r values P value correlation P value
Matching
Cranial metric and nonmetric 0.1140 0.1810 0.6626 0.0260
Cranial metric, Y-chromosome 20.2054 0.6170 0.6982 0.5230
Cranial metric, mtDNA 0.6812 0.0080 0.7668 0.0090
Cranial nonmetric, Y-chromosome 0.7626 0.0240 0.9095 0.0290
Cranial nonmetric, mtDNA 20.2915 0.5090 0.4681 0.3860
Unmatching
Cranial metric and nonmetric 0.7896 0.0800 0.8234 0.0540
Cranial metric, Y-chromosome 0.8000 0.0770 0.8668 0.0520
Cranial metric, mtDNA 0.3506 0.2640 0.8792 0.2060
Cranial nonmetric, Y-chromosome 0.7645 0.0670 0.6792 0.1990
Cranial nonmetric, mtDNA 0.4680 0.1600 0.8641 0.3540

Bold indicates significance at the 0.05 level.

Fig. 2. Graph of procrustes analysis using matching metric and nonmetric data.

Fig. 3. Graph of procrustes analysis using matching metric and mtDNA data.

algorithms to converge. Thus, only cranial phenotypic
data will be considered. Two-tailed P values for the Man-
tel test were generated, using multiple combinations of
the data, to test the null hypothesis that these popula-
tions are not genetically related. The two-tailed P values
are listed in Table 8 (all R matrices provided in the Sup-
plementary Information). A two-tailed P value is more
appropriate to use in this case, rather than the one-
tailed P value, because the directionality of the relation-
ship between the different data sets is not limited to one
side or the other. The null hypothesis for the Mantel test
is that the data sets are not correlated. Using this level

American Journal of Physical Anthropology

8 B. HERRERA ET AL.

Fig. 4. Graph of procrustes analysis using matching metric and Y-chromosome DNA.

Fig. 5. Graph of procrustes analysis using matching nonmetric and mtDNA.

Fig. 6. Graph of procrustes analysis using matching nonmetric and Y-chromosome DNA.

of significance, two out of the eleven combinations are
statistically similar (the matching metric and mtDNA,
and matching nonmetric and Y-chromosome), allowing
us to reject the null hypothesis for these two combina-
tions of data. The PROTEST, which is a better determi-
nation of the underlying relationship between data sets,
demonstrates statistically significant similarities
between an additional combination (the matching metric
and nonmetric set). The mtDNA and Y-chromosome
DNA were not run against each other because it would
be a correlation between two different sets of inheri-
tance. Except in the case of inbreeding, any correlations
might be spurious.
The results of the Procrustes analysis were graphed
using the Scatterplot3D package (Ligges and M€ achler,
2003) in R, shown in Figures 2–12. The matching sam-
ples are shown in Figures 2–6. Figure 2 shows the Aleut
samples grouping separately from the other populations.
In general, they match most closely with the nonmetric
Canada, Alaska 1 (Alaskan Yupik), and Alaska 2

American Journal of Physical Anthropology

 COMPARABILITY OF MULTIPLE DATA TYPES 9

Fig. 7. Graph of procrustes analysis using unmatching metric and nonmetric data.

Fig. 8. Graph of procrustes analysis using unmatching metric and mtDNA.

Fig. 9. Graph of procrustes analysis using unmatching metric and Y-chromosome DNA.

(Alaskan Inupiat) populations. All samples appear to the Aleut samples grouping together, with the exception
match with their counterpart in all three dimensions— of the mtDNA Aleut 3 (Bering Island), which is closest
principal coordinates 1, 2, and 3. Figure 3 also shows to the mtDNA Siberia sample (Chukchi).

American Journal of Physical Anthropology

10 B. HERRERA ET AL.
Fig. 10. Graph of procrustes analysis using unmatching nonmetric and mtDNA.
Fig. 11. Graph of procrustes analysis using unmatching nonmetric and Y-chromosome DNA.
Focusing on only the statistically significant combina-
tions, the matching metric and mtDNA Procrustes anal-
ysis show a clear distinction between the Aleutians and
the rest of the populations, except the Aleut three group.
The same is true for the matching metric and nonmetric
combination, with a similarity in location of the points
in all dimensions from all populations, except in the case
of the Canadian population. The matching nonmetric
and Y-chromosome set has very similar locations
between all points in all dimensions, with the farthest
apart being the Aleut 1 population.
Most of the samples are similar to their counterpart in
all three dimensions, with the exception of Alaska 1,
Alaska 2, and Aleut 6. Figure 4 does not strongly show
this three-dimensional similarity with the samples used.
This represents the higher P values from Table 8. The
Aleut 1 metric sample groups away from the others, as
does the Greenland metric sample. Similarly, the Aleut 1
Y-chromosome sample is closest to the Alaska 1 Y-
chromosome sample (Yakut). The Greenland Y-
chromosome sample (Inuit) is closest to the Alaska 2
(Inupiat) metric sample. Figure 5 shows a stronger
grouping of data type samples instead of population
samples. The nonmetric samples have several groupings:
Aleut 4 and Canada, Aleut 6 and Siberia, Alaska 2 and
Greenland, and finally Aleut 2, Alaska 1, and Aleut 1.
American Journal of Physical Anthropology
The mtDNA groupings are as follows: Aleut 1, 3, 4, fol-
lowed by Alaska 1 and Aleut 2, and Alaska 2, Green-
land, Aleut 6, Canada, and Siberia. Figure 6
demonstrates similarity in all dimensions between all
sample combinations, but the Aleut 1 nonmetric sample
(Unalaska) matches most closely with the Alaska 1 Y-
chromosome sample (Western Alaskan Yakut). The oppo-
site is also true, with the Aleut 1 Y-chromosome sample
(Unalaska) matching most closely with the Alaska 1
nonmetric sample (Yakut).
The unmatching Procrustes graphs are provided in
Figures 7–11. Figure 7 demonstrates similarity in all
dimensions for all population pairs. The exception to
this is that there is a large difference in principal coordi-
nates 2 and 3 for the Alaska 2 population pair. The
Aleut populations cluster separately, but nearby to the
other populations. The Greenland population pair is the
farthest removed. Figure 8 has similarities between all
dimensions for the population pairs. The Aleutian popu-
lations group separately, as do the two Alaskan popula-
tion pairs and the Greenland population pair. The same
holds true for Figure 9. In Figure 10, the Aleut 1 and
Alaska 1 population pairs group together. Otherwise, the
mtDNA Alaska 2 (Inupiat) and the mtDNA Greenland
populations group together. Lastly, Figure 11 shows mul-
tiple groupings: nonmetric Alaska 1 (Yupik) and 2
 COMPARABILITY OF MULTIPLE DATA TYPES
(Inupiat) with Y-chromosome Greenland, as well as Y-
chromosome Alaska 1 (Yupik) and 2 (Inupiat) with non-
metric Greenland Inuit. The Aleut populations group
separately and match in three-dimensions except for
principal coordinates 2 and 3 for Aleut 1.
DISCUSSION
P values from Mantel tests of the matching data indi-
cate that the craniometric and mtDNA data combination
have a statistically significant correlation. The same
holds true for the cranial nonmetric and Y-chromosome
combination. More simply stated, the two types of data
provide similar information. These significant correla-
tions extend beyond the phenotype to the genetics; the
two phenotypes (metric and nonmetric) also demonstrate
a significant association between them. The concordance
of the skeletal data to the molecular data will be
described in detail, below, as well as population affinities
of the Northern demes and application of this methodol-
ogy to peopling of the New World hypothesis testing.
The P values associated with the PROTEST analysis
indicate that three combinations (matching cranial met-
ric and nonmetric, matching cranial metric and mtDNA,
matching cranial nonmetric and Y-chromosome) all can
reject the null hypothesis at the 0.05 significance level.
Two additional combinations (the unmatching cranial
metric and nonmetric, and the unmatching metric and
Y-chromosome) are equivalent to 0.05 and still represent
a significant relationship among the data types. These
results are consistent with work by Ricaut et al. (2010)
who found similar correlations between nonmetric data
and genetic markers, demonstrating some degree of her-
itability with nonmetric traits. Other studies have also
found some degree of heritability in cranial metric or
nonmetric traits (e.g., Sjïvold, 1984; Cheverud, 1988;
Relethford, 1994; Sparks and Jantz, 2002; Carson,
2006a, 2006b; Martınez-Abadıas et al., 2009).
The Mantel and PROTEST results support the neutral
theory of evolution, where genetic and phenotypic dis-
tances would be similar or proportional (Roseman and
Weaver, 2007). This also supports Martınez-Abadıas
et al. (2009), in that our mostly large sample sizes may
have circumvented the issues they noted with discon-
cordance in smaller sample sizes. The significant associ-
ation between the unmatched cranial metric and Y-
chromosome data could be spurious, which is supported
by the lack of significance in the matched cranial metric
to Y-chromosome data. However, the sample size and
demes differed between the matched and unmatched
data sets, which may have led to a difference in signifi-
cance between the two. With these conflicting results,
the most appropriate interpretation is the conservative
approach: the association between craniometrics and Y-
chromosome data is inconclusive. The confirmed associa-
tion between the specific molecular and phenotypic data
may be as a result of similar evolutionary forces. It
appears to be the case that evolutionary forces affecting
the Y-chromosome also affect nonmetric cranial traits.
The same may hold true for metric traits and mtDNA.
The Procrustes graphs generally reflect the P values
shown in Table 8, with the statistically significant com-
binations showing more concordance in the graphs.
These graphs clearly show a distinction between the
Aleut and other populations. The demes with the most
similarity to the Aleutian ones were, in general, Alaska
1 and 2. Most of the demes grouped according to their
11
counterpart in the corresponding data type. This holds
particularly true for the combinations that were found
to be statistically significant (see Table 8).
For the most part, the samples clustered by population
affinity across data types, supporting the respective pop-
ulation histories. In general, the graphical differences
among correlated data types appear to be complimen-
tary, rather than competing. Thus, when combined they
provide a more complete picture of population relation-
ships. The Aleutian and Alaskan samples typically
appeared nearby in the figures, which is consistent with
documented cultural diffusion (Corbett et al., 1997). Spe-
cifically, Figures 2 and 5 show Aleut2 clustered near the
Alaskan demes. This Aleut sample is comprised of indi-
viduals from east of the Rat Islands. Cultural diffusion
is evident all the way to the Rat Islands and the Near
Islands, where the level of shared culture decreases dra-
matically (Corbett et al., 1997). Thus, the relationship of
Alaskan Inuits with Aleutians east of the Rat Islands
makes sense in light of the ethnographic and archaeolog-
ical evidence. Additionally, Greenland and Alaskan
demes clustered together in Figures 2–5 and 8–11.
The Alaskan Yupik, Inupiat, and Greenland Inuit rela-
tionship can be confirmed through genetic evidence.
There is currently debate about the exact relationship
between the Alaskan Thule (ancestors to both the Alas-
kan Yupik and Inupiat) and the Greenlandic Inuit (Sail-
lard et al., 2000; Melgard et al., 2005; Dulik et al.,
2012). This debate centers around whether the Thule
alone gave rise to the Greenlandic Inuit, or whether the
Inuit formed from interactions between the Dorset and
Thule (Dulik et al., 2012). In either case, there is evi-
dence for gene flow between the Thule and Inuit. There
is some historical evidence as well. The Thule adopted
several Dorset technologies, such as snow houses and
knives (Stern, 2010). It is also known that the Norse
had interactions with the Thule and Inuit in Greenland
(Stern, 2010). These historical and genetic relationships
are again uncovered by our combination of Y-
chromosome and cranial nonmetric data.
No skeletal data type should display a perfect correla-
tion with DNA; environmental influence prohibits this
from occurring. What is interesting in the findings here
is some correlations with genetic data are high, indicat-
ing some heritability of these traits when considered as
a group. The amount of probable heritability lends sup-
port for using cranial metrics or nonmetrics as proxies
for genetic material. Specifically, this work shows that
cranial metrics can likely be used as a proxy for mtDNA
haplogroup frequencies and cranial nonmetrics can
likely be used as a proxy for Y-chromosome DNA fre-
quencies. Moreover, craniometrics and cranial nonmet-
rics agree, here, showing that one is not necessarily
superior to the other.
The evidence that some heritability exists (although
not quantified) amongst the cranial metric and nonmet-
ric traits is important for researchers wishing to investi-
gate greater bioarchaeological hypotheses, in particular,
peopling of the New World. Peopling of the New World is
mentioned here because of the lack of agreement
between data types resulting in conflicting hypotheses.
Ancient DNA is difficult to obtain from important skele-
tal specimens for peopling of the New World, dating
back 15000–20000 BP. Therefore, furthering the under-
standing of the complicated relationship between cranial
data and genetics is important for improving upon cur-
rent methodologies. Further, while there are seemingly
American Journal of Physical Anthropology
12 B. HERRERA ET AL.
contradictory studies on peopling of the New World
involving either cranial morphology or genetics (Green-
berg et al., 1986; Bonatto and Salzano, 1997; Powell and
Neves, 1999; Schurr and Sherry, 2004; Tamm et al.,
2007; Mulligan et al., 2008; Perez et al., 2009; Hubbe
et al., 2010; Pitblado, 2011), it is likely that each of these
data types are showing only a portion of the whole story,
as evidenced in the figures.
Our work is not without bias. Although, the mtDNA/
craniometric and Y-chromosome/cranial nonmetric pat-
terns emerged more than once, indicating we are prop-
erly modeling the structure of this data set, we cannot
discount that these findings are population specific. Fur-
ther, our samples were all from the same time period
(all contemporary), but the exact census population sizes
are not known. The same people within each population
also could not be used to obtain both DNA and cranial
metric/nonmetric traits, which could have introduced
error and implies that the correlations are only the mini-
mum estimate (Roseman, 2004). Without further studies
involving extraction of all data types from the same indi-
viduals, the extent to which this affected the results
shown are unknown. To best remedy this potential bias,
we used similar demes to compare to each other. Each
group used to approximate another has a known shared
history or interaction with the other. This increased the
odds that there was gene flow between these peoples,
and thus the demes should reflect similar genetic
variation.
In sum, the results here demonstrate that cranial phe-
notypic data (metric and nonmetric) are linked and are
excellent proxies for mtDNA and Y-chromosome DNA.
The correlations of phenotypic and genetic data are
highly specific (mtDNA to metrics and Y-chromosome to
nonmetric). Although exact heritabilities were not calcu-
lated here, the high concordance of the molecular and
phenotypic data indicates that some genetic heritability
is present among these data types. The biological affin-
ities uncovered were consistent with population histor-
ies, indicating our methodology is solid. Our study does
not utilize temporal and spatial components, so this
study alone is not sufficient for interpretations of the
human migration into the New World. In spite of this,
our study has implications for research relating to
migration. Knowing how well each data type compli-
ments each other is important for interpreting the vast
amount of information different lines of research pro-
vide. More inclusive research involving various data
types and disciplines needs to be completed so a better
understanding of human events, e.g., migration, can
occur.
ACKNOWLEDGMENT
The authors thank Dr. Mark Hubbe, an Associate Edi-
tor for AJPA, and the anonymous reviewers for their
comments and suggestions.
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