Intragroup variation in the Pre-Columbian Cuba population: A perspective from cranial morphology

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Intragroup variation in the Pre-Columbian Cuba population: A perspective from cranial morphology
Pre-Columbian Cuba population cranial morphology

                                                             Taisiya Syutkina et al.

                                     Anthropological Review • Vol. 84(3), 233–255 (2021)

          Intragroup variation in the Pre-Columbian
         Cuba population: A perspective from cranial
                         morphology

                     Taisiya Syutkina1, Mario Juan Gordillo Pérez2,
            Silvia Teresita Hernández Godoy3,4, Carlos Arredondo Antúnez5,
                                Armando Rangel Rivero6
     1
       Center for Human Ecology, the Russian Academy of Sciences N.N. Miklouho-Maklay
                           Institute of Ethnology and Anthropology
                2
                  Department of Biology, University of Oriente - Santiago de Cuba
    3
      Grupo de Investigación y Desarrollo de la Dirección Provincial de Cultura de Matanzas
             4
               Departamento de Estudios Socioculturales, Universidad de Matanzas
         5
           Montané Anthropological Museum, Biological Faculty, University of Havana
         6
           Montané Anthropological Museum, Biological Faculty, University of Havana

Abstract: The paper aims to study intragroup variation inside the two pre-Columbian Cuban populations:
the aceramic Archaic and the ceramic Taino groups, based on their cranial morphology. The latter applied
artificial cranial deformation to all its members, so the groups are referred to as “non-deformed” and
“deformed” samples here. Studies across different disciplines suggest evidence of cultural and biological
diversity inside the non-deformed group, while local variations of applying the deforming device can be
responsible for shape variation across the deformed group. Cranial metrics and non-metric cranial traits of
the 92 crania of Cuban origin were analyzed, although the sample size varied between the analyses due to
the incompleteness of the crania. Geometric morphometrics was applied to the deformed crania to study
the shape variation across the sample. Three deformed crania from the Dominican Republic were ana-
lyzed together with the deformed Cuban sample to test the variability of the practice between the islands.
Principal component analysis and the Mantel test did not reveal any geographic differences in the cranial
metric traits. No morphological differences associated with the antiquity of materials could be seen either
based on the available data. The principal component analysis of the Procrustes coordinates of the cranial
vault outline in the lateral norm revealed continuous variability of cranial shapes from the ones with more
flattened frontal and occipital bones to the more curved outlines, which is probably explained by individual
variation. Non-metric traits variation revealed bilateral asymmetry in the expression of the occipito-mas-
toidal ossicles among the deformed crania. In conclusion, the study did not support assumptions about
morphological diversity inside the studied samples or proved the impossibility of available craniological
data to reflect possible intragroup differentiation at the moment.
Key words: biological anthropology, cranial metrics, non-metric traits, geometric morphometrics, anthro-
pology of Cuba, artificial cranial deformation, The Greater Antilles

Original Research Article Received: June 25, 2021; Revised: August 23, 2021; Accepted: August 27, 2021
DOI: 10.2478/anre-2021-0021
© 2021 Polish Anthropological Society
Intragroup variation in the Pre-Columbian Cuba population: A perspective from cranial morphology
234                                Taisiya Syutkina et al.

             Introduction                       ticular type requires both deformed and
                                                non-deformed crania from the same pop-
Prior to European arrival, Cuba is known        ulation, and the results are only valid for
to have been inhabited by an indigenous         the population in question (Rhode and
population, which is generally divided          Arriaza 2006). Possible intragroup varia-
into two groups: the aceramic Archaic           tion can be explained by multiple origins,
population, which is believed to have ar-       microevolutionary processes, different
rived around 5000 years ago (Napolitano         lifestyles for the non-deformed sample
et al. 2019:7–8) and the ceramic agricul-       and by local traditions in applying the
tural Taino population, which inhabited         deforming apparatus for the deformed
the island since approximately 500 AD           sample. Apart from the spatial dimen-
(Torres Etayo 2006:35–36) until deci-           sion, there is a temporal one, which
mation by the newcomers from the Old            could reflect possible changes through-
World.                                          out the long history of the island popu-
    There has been much variation in the        lation. The assumption that these factors
ways the aboriginal population of Cuba          might have impacted the morphology of
is referred to. To avoid confusion caused       the studied groups follows from the re-
by changing perspectives due to new ar-         sults of numerous studies across differ-
chaeological, linguistic and biological ev-     ent disciplines.
idence, we will stick to strictly morpho-           Two recent genetic studies of the
logical criteria of “non-deformed” and          pre-contact Caribbean populations, in-
“deformed” crania, which are associated         cluding the Cuban ones (Nägele et al.
with the aceramic (Archaic) and ceramic         2020; Fernandes et al. 2021), confirmed
(Taino) population groups respectively.         a single genetic profile for Ceramic-age
    This study focuses on the intragroup        (referred to as “deformed” in this study)
variation of the Cuban pre-Columbian            individuals, which suggests a single mi-
population basing on cranial morphol-           gration or successive migrations from
ogy of the deformed and non-deformed            the same region for this group. Howev-
samples. Between-group variation is not         er, the studies produced different results
being addressed, and the samples are an-        considering the homogeneity of the Ar-
alyzed separately for two reasons. First,       chaic (“non-deformed”) population. This
there has been extensive evidence that          inconsistency is most likely explained by
the two groups did not have recent com-         different sampling: the Cueva del Perico
mon genetic ancestry, so comparing their        individual, shown to have different (pre-
cranial morphologies would not yield            sumably North American) ancestry by
new information. Second, a large body           the work of Nägele et al., was not includ-
of evidence suggests that artificial cranial    ed in the second genetic study, neither
deformation affects facial and basal areas      was it available for the present work.
of the cranium (Antón 1989; Cheverud                An extensive odothological study has
et al. 1992; Friess and Baylac 2003), so        identified two migratory waves that led
the direct comparison of deformed and           to populating the Circum-Caribbean re-
non-deformed crania of different popula-        gion, one connected with the aceramic
tions would have produced spurious re-          Archaic groups, the other one bringing
sults. Detection of the regions that stay       the ceramic Taíno population (Coppa et
unaffected by the deformation of a par-         al. 2008). The deformed Cuban sample
Intragroup variation in the Pre-Columbian Cuba population: A perspective from cranial morphology
Pre-Columbian Cuba population cranial morphology                235

in the study was represented by the sam-       to a more differentiated approach to the
ple coming from a single cemetery, Chor-       diversity of pre-Columbian communities
ro de Maíta, while the non-deformed            in Cuba.
samples came from multiple sites and               Archaeological data also suggested a
still clustered together according to their    possible division of the non-deformed
dental morphology. This finding is espe-       population into two periods: hunt-
cially remarkable as one of the samples        er-gathering and protoagricultural (ac-
analyzed by Coppa et al. came from Cue-        tual names may vary from author to au-
va del Perico, an outlier in the genetic       thor), the latter one characterized by the
study discussed above.                         evidence of simple agriculture and crude
    Archaeological data suggest complex        ceramics at the corresponding sites
population interconnections within the         (Tabío 1988:64–65; Guarch Delmon-
island at different periods (Cooper 2007;      te 1990; Pérez Carratalá 2014:78–80).
Chinique de Armas et al. 2020), while,         It has also been shown based on stable
on the other hand, some sites stayed           isotope analysis that subsistence strat-
isolated from the surrounding groups           egies differed throughout the territory
throughout long periods. For example,          of the island even between contempora-
Canímar Abajo in Matanzas (populat-            neous communities: groups inhabiting
ed by the non-deformed group) bears            Canímar Abajo site have been shown to
evidence of long-standing cultural tra-        rely on horticulture while Cueva Calero
ditions not present at other contempo-         and Cueva del Perico inhabitants appar-
raneous Cuban sites (Alarie and Rock-          ently relied on hunting-fishing-gather-
sandic 2016). This long isolation could        ing strategy (Chinique de Armas et al.
have formed the basis for possible mi-         2018).
croevolutionary processes within some              Classification of the archaeological
communities, which might have affected         and skeletal materials is also an essential
their cranial morphology.                      issue for discussing the groups’ homo- or
    Previously the non-deformed group          heterogeneity. Cooper (2007:232) noted
was often divided into two archaeologi-        that ceramic vessel fragments had been
cal complexes (Herrera Fritot 1964; Po-        found at some sites classified as pre-agri-
spisil and Rivero de la Calle 1964; Tabio      cultural in the Western part of the island,
and Rey 1966): Complex I and Complex           and the cultural attribution relied upon
II (sometimes called as Guayabo Blan-          the absence of artificial cranial deforma-
co and Cayo Redondo or Guanahatabey            tion in the human remains. He argues
and Siboney). The division was based on        that it challenges the whole system of
mortuary practices registered at two dif-      classification of the findings, which au-
ferent sites, but some researchers tend-       tomatically considers any non-deformed
ed to see two biologically distinct pop-       crania belonging to the pre-agricultural
ulations behind them (Alonso Alonso            population. This, in turn, means that one
1995:99). Later a more profound consid-        has no reason to assume that all non-de-
eration of ancient cultures made for un-       formed crania belonged to a morpholog-
derstanding that the interplay between         ically, genetically, or culturally uniform
the groups that occupied different sites       population.
was more complex than a simple dichot-             As far as the deformed sample is con-
omy of two cultural variants, which led        cerned, Herrera Fritot (1964:105) noted
Intragroup variation in the Pre-Columbian Cuba population: A perspective from cranial morphology
236                                Taisiya Syutkina et al.

that the deformed crania could be divid-            Cranial samples from Cuba avail-
ed into two groups: those with moder-           able for this study suffer from several
ate deformation concentrated mainly in          problems: most of them lack contextu-
the frontal and basal occipital areas and       al information, zone of origin is often
those with more pronounced deforma-             known approximately, secure dating is
tion which affected the whole vault and         only available for several sites. Moreover,
in most cases produced a bilobate shape.        the humid climate does not favor the
He deduced that there must have existed         long conservation of the bone remains,
different designs of deforming devices,         so their state of preservation is often far
some of them having a sagittal band in          from perfect. With that in mind, this ar-
addition to the transverse ones.                ticle addresses the following questions:
    The results of all the studies men-         (1) Did possible multiple origins or mi-
tioned above suggest grounds for ex-            croevolutionary processes contribute to
pecting a certain degree of heterogeneity       morphologically detectable intragroup
both inside the deformed and non-de-            differentiation inside the non-deformed
formed samples. There is evidence that          sample? (2) Did local differences in ar-
some local non-deformed groups relied           tificial cranial deformation contribute
upon different subsistence strategies,          to intragroup differentiation inside the
were culturally diverse and had lived in        deformed sample? and (3) Can analysis
isolation for a long time (Chinique de          of the available craniological data con-
Armas and Rocksandic 2019). Consid-             tribute to our understanding of the rela-
ering that they had been inhabiting the         tionships between different groups that
island for several millennia, these factors     inhabited the island before European
might have led to significant differences       colonization?
between the local or temporal groups.               Cranial morphology is widely used
Given that the non-deformed sample can          to address prehistorical issues due to
include individuals with different ances-       the conservative nature of its variation,
try and that there is no certainty about        its underlying genetic component, and
them being a single group, a study of           strong geographic patterning (Bunak
intragroup morphological variability can        1959; Howells 1989; Pietrusewsky 2014).
provide insights into the anthropological       Metric and non-metric cranial traits were
structure of the island’s early popula-         used in this study to explore the variabil-
tion. While not much genetic-based in-          ity of cranial morphology in each sample.
tragroup morphological variation can be         For the deformed group, two-dimension-
expected for the deformed sample, little        al geometric morphometrics was applied
is known about the deforming practice           to study and visualize the shape variation
itself and whether it was performed in          across the sample. 2D shape analysis is
the same manner throughout the whole            a robust tool for studying morphologi-
island. The objective of the present study      cal variation but must be used with cau-
is to analyze the intragroup variation in       tion with 3D objects due to the so-called
cranial morphology of the two pre-Co-           “three-to-two-dimension error” (Cardini
lumbian Cuban samples and see wheth-            2014; Buser et al. 2018), and its perfor-
er individual differentiation inside them       mance has been shown to deteriorate
correlates with territorial, cultural, or       when applied to small samples belong-
temporal factors.                               ing to one taxon (Cardini and Chiapelli
Pre-Columbian Cuba population cranial morphology                237

2020). However, it has been proven ef-          de la Calle (1964) questioned this idea:
fective for the studies of artificial cranial   according to them, it was difficult to sep-
deformation (Perez 2007; Gonzalez et al.        arate the first two complexes both by ar-
2011) and thus will be only applied to          chaeological and anthropological data. A
the deformed group in this paper.               direct comparison between the crania did
                                                not reveal any differences, so it was sug-
  A review of previous cranial                  gested that the preceding authors might
                                                have come to an opposite conclusion due
   studies of the Cuban pre-                    to the inclusion of both male and female
     Columbian population                       crania in their samples.
                                                    Soviet anthropologist V. Ginzburg,
Various scholars attempted to study the         whose measurements of the Cuban cra-
internal structure of the Cuban pre-Co-         nia were included in the present paper,
lumbian population basing on cranial            concluded that the non-deformed and
data, which has always been scarce. The         deformed groups were not directly re-
first scholar to study ancient human re-        lated, although both must have had
mains was Luis Montané Dardé, who               South-American biological affinities. He
studied the non-deformed crania from            also suggested two morphological sub-
the site of Boca del Purial and noted their     types of the non-deformed crania, one
anthropological heterogeneity (Mon-             being wider- and lower-faced than the
tané 1908 cited in Hernández Godoy              other (Ginzburg 1967:189). His conclu-
2003:192). Later studies were conducted         sions were disputed by V. Alexeev, who
by Rivero de la Calle, who examined the         suggested that the differences between
deformed crania and identified this type        the groups could be entirely explained
of deformation as “tabular oblique” ac-         by the effect of cranial deformation
cording to Dembo and Imbelloni’s clas-          (Alexeev 1986:20). By studying the vari-
sification (1938) and detected one crani-       ation of non-metric traits in the two pop-
um that had “tabular erect” type, which         ulations, Rivero de la Calle (2009:179–
he explained as a possible defect of ap-        180) concluded that their divergence is
plication of the deforming device (1949         greater than the one between compara-
cited in Herrera Fritot 1964:82–83).            tive samples from distant locations of
    One of the first works summarizing all      the American continent (like Peru and
the known cranial data was made by René         British Columbia). Recent genetic stud-
Herrera Fritot (1964), who developed            ies reaffirmed that both groups had se-
the craniotrigonometric method and ap-          curely different origins (Lalueza-Fox et
plied it to the study of Cuban aboriginal       al. 2003; Nägele et al. 2020; Fernandes
crania. He investigated them through the        et al. 2021).
prism of archaeological classification of           After that, the focus of major archae-
the three archaeological “complexes”–           ological and anthropological works based
Complex I or Guayabo Blanco (non-de-            on Cuban pre-Columbian materials
formed), Complex II or Cayo Redondo             shifted from studying the population in
(non-deformed), and Complex III or              general to studying local cultural, ecolog-
Taíno (deformed), and saw individuals           ical, nutritional, health, and other issues
from the respective sites as characteris-       (Crespo-Torres et al. 2013:437–439).
tic for these “types”. Pospisil and Rivero      This turned to be a promising approach;
238                                Taisiya Syutkina et al.

biological anthropology managed to shed         the pre-Columbian Cuban population us-
new light on mortuary (La Rosa Cor-             ing a multimethodological approach and
zo 2003), weaning (Chiniqie de Armas            taking into account findings reported in
2017) and cultural practices (Alarie and        recent studies from other disciplines and
Roksandic 2016), subsistence strategies         new perspectives on the complex rela-
(Chinique de Armas et al. 2015), nature         tions of the groups inhabiting the island.
of co-existence with the Europeans at
the later sites (Valcárcel Rojas 2016), pa-             Material and methods
thologies (Armstrong et al. 2013).
    Some recent attempts to approach in-        Data acquisition. The total craniometric
tragroup diversity based on morpholog-          sample consists of 95 crania: 92 male and
ical data were proposed by students of          female crania of Cuban origin and three
The University of Havana. Bolufé Torres         deformed male crania from the Domin-
(2015) found differences in the orbital         ican Republic to test the variability of
breadth and stature between non-de-             cranial morphology between the islands
formed individuals from different loca-         (see Table 1 for more information on the
tions in the West of the island. Although       sites and Supplement Table 1 for the full
the author concluded that the results           composition of the samples). The Cuban
might indicate the different origin of the      sample is comprised of the 57 non-de-
individuals, they should be treated with        formed (male = 40, female = 17) and
caution due to the small sample sizes           39 deformed (male = 27, female = 12)
available. Valdés Pi (2009) managed to          crania. However, not all of them could
compose a comparatively large sample of         be included in all the multivariate anal-
both non-deformed and deformed crania.          yses due to their incompleteness. The
A set of cluster analyses based on facial       sample size for each analysis is reported
craniometric variables identified possi-        separately in this section. Cranial mea-
ble sub-clusters both inside the non-de-        surements have been standardized by
formed and deformed groups, but no              both the Biometrika school and Martin
patterns (geographical, cultural, chrono-       and Saller (1957) in the version applied
logical) that could explain the identified      in Soviet/Russian craniometry (Alexeev
division were suggested. García Méndez          and Debets 1964). The craniometric
carried out a geometric morphometric            sample is composed of two parts: crania
analysis of the outlines characterizing         measured by Vulf Ginzburg (n = 18) in
the cranial shapes in the frontal and lat-      1964 (Ginzburg 1967) and crania mea-
eral norms in a sample of non-deformed          sured by one of the authors (TS) in
individuals from the East and the West          2017 following the same methodology
of the country. She concluded there were        (n = 77). Ginzburg’s sample consists
no significant differences between them         of the crania that could not be found or
(García 2018; García et al. in press). Fi-      accessed in 2017 and only contains cra-
nally, preliminary analyses of the metric       niometric variables. Non-metric traits
data studied here have been previously          were observed by TS and MGP for the
published in Russian by the first author        same 77 individuals that were measured
(Syutkina 2017; 2018).                          by TS. The traits used in the study are a
    This paper is the first attempt to an-      combined list of traits described in Berry
alyze a comprehensive cranial sample of         and Berry (1967), Krenzer (2006), and
Pre-Columbian Cuba population cranial morphology                            239

Fig. 1. Map showing location of the sites discussed in the article. Generated with the use of raster package
    in R (Hijmans 2021). The sites from the zone of Baracoa are represented as a single point, more infor-
    mation on exact origins is available in the Supplement Table 1

Table 1. Cranial samples from Cuba and Dominican Republic
                                     Site                                Male       Female         Total
Non-deformed Unknown                                                       2           0              2
             Baracoa                                                      16           7            23
             Punta del Este                                                1           0              1
             Pinar del Río                                                 1           0              1
             Cueva Calero, Matanzas                                        0           3              3
             Boca del Purial, Sancti Spíritus                              1           3              4
             Canímar Abajo (Younger Cemetery), Matanzas                   10           1            11
             La Santa, Habana                                              0           1              1
             Cueva Florencio, Matanzas                                     1           0              1
             Ciénaga de Zapata, Matanzas                                   2           0              2
             Guayabo Blanco, Matanzas                                      2           0              2
             Puerto de Santa María, Camagüey                               3           1              4
             Laguna del Tesoro, Matanzas                                   0           1              1
             Total non-deformed                                           39          17            56
Deformed     Unknown                                                       3           3              6
             Baracoa                                                      18           6            24
             Guardalavaca                                                  1           0              1
             Cayo Salinas, Sancti Spíritus                                 2           1              3
             Banes                                                         0           2              2
             Cueva de Yasica, Dominican Republic                           1           0              1
             La Caleta, Dominican Republic                                 2           0              2
             Total deformed                                               27          12            39
240                                   Taisiya Syutkina et al.

Movsesyan (2005) (see Table 2 for the              era, skulls placed in lateral view, camera
full list and references). All traits were         positioned at 30 cm from the auriculare
registered per side to study the symme-            point. Two landmarks (the deepest point
try of their expression and per cranium            on the frontal bone after glabella and the
to calculate their frequencies in each             intersection of the line continuing the di-
group (Czarnetzki 1971). Only adult cra-           rection of the zygomatic process of the
nia were included (based on the fusion of          temporal bone with the outline of the
the spheno-occipital synchondrosis). Sex           skull in its posterior part) and 23 semi-
and age for the 77 crania were estimat-            landmarks between them were placed
ed according to general methodological             along the sagittal curve (Figure 2). These
guidelines (Buikstra and Ubelaker 1994).           landmarks were chosen to analyze the
    The shape of the deformed crania               outline avoiding the glabella and mastoid
in the lateral norm was studied using              regions, which would otherwise add un-
two-dimensional geometric morpho-                  desirable sex-related variation. The land-
metrics, following standard procedures             marks were placed, and their coordinates
(Bookstein 1991; Zelditch and Swiderski            were obtained using tpsDIG2 2.31 soft-
2018; Vasiliev et al. 2018). Images were           ware (Rohlf 2015). The semilandmarks
taken with a Nikon Coolpix P100 cam-               were then slid according to the minimum

Table 2. Non-metric traits used in the study and their frequencies in the non-deformed and deformed
    samples
              Trait                      Reference         Non-deformed n (N) Deformed n (N)
Metopism                           Berry and Berry 1967          0 (40)           0 (33)
Saggital ossicles                      Krenzer 2006              0 (29)         0.129 (31)
Ossicle at the lambda              Berry and Berry 1967        0.071 (28)       0.188 (32)
Inca bone                              Krenzer 2006              0 (28)         0.125 (32)
Palatine torus                     Berry and Berry 1967          0 (33)         0.103 (29)
Supraorbital foramen                   Krenzer 2006            0.538 (39)       0.406 (32)
Supraorbital notch                     Krenzer 2006            0.795 (39)       0.906 (32)
Accessory infraorbital foramen     Berry and Berry 1967        0.216 (37)       0.133 (30)
Infraorbital suture                    Krenzer 2006            0.649 (37)       0.600 (30)
Trochlear spine                      Movsesyan 2005            0.091 (33)       0.000 (28)
Multiple zygomatic foramina            Krenzer 2006            0.784 (37)       0.607 (28)
Stellate pterion               Movsesyan 2005, Murphy 1956     0.029 (35)       0.034 (29)
Fronto-temporal articulation       Berry and Berry 1967        0.000 (35)       0.034 (29)
Epipteric bone                     Berry and Berry 1967        0.000 (35)       0.207 (29)
Coronal ossicles                   Berry and Berry 1967        0.057 (35)       0.032 (31)
Lambdoid ossicles                  Berry and Berry 1967        0.148 (27)       0.533 (30)
Occipito-mastoidal ossicles            Krenzer 2006            0.348 (23)       0.533 (30)
Ossicle at the asterion            Berry and Berry 1967        0.074 (27)       0.367 (30)
Squamoparietal ossicles                Krenzer 2006            0.032 (31)       0.414 (29)
Parietal foramen                   Berry and Berry 1967        0.767 (30)       0.793 (29)
Foramen of Huschke                 Berry and Berry 1967        0.414 (24)       0.310 (29)
Posterior condylar canal           Berry and Berry 1967        0.962 (26)       0.880 (25)
Condylar facet double              Berry and Berry 1967        0.188 (16)         0 (20)
Pre-Columbian Cuba population cranial morphology                 241

                                                male and female samples and compared
                                                with average world figures (Alekseev and
                                                Debets 1964:123–25) where available.
                                                SSD was only calculated for traits that
                                                could be measured in no less than ten
                                                individuals in the smaller female sample
                                                (Evteev 2008:9).
                                                    Principal component analysis was
                                                performed to study the patterns of mor-
                                                phological variation within each sam-
                                                ple. Metric variables were used for the
                                                non-deformed sample: the set of variables
Fig. 2. Landmarks (squares) and semilandmarks
                                                to be included was composed in a manner
    (circles) used in the study                 that would allow maximizing the sample
                                                size and cumulative proportion of the
bending energy criterion, and both land-        variance explained by the first two PCs.
marks and semilandmarks were aligned            For the same reason, neurocranial and
using a generalized Procrustes analysis         facial variables were studied separately
(Bookstein 1991).                               for the non-deformed sample, as com-
                                                bining them led to a drastic decrease of
           Statistical analyses                 the sample size to a very small number of
                                                individuals. Seven facial measurements,
All craniometric values were compared           summarizing overall facial, nasal, and or-
with the worldwide variation of the             bital dimensions (FMB, ZMB, UFH, NLB,
studied traits and categorized as “very         NLH, OBH, OBB) and ten measurements
small”, “small”, “medium’, “large”,             describing cranial vault (GOL, XCB, BBH,
“very large” (Alexeev and Debets                AUB, FRA, PAA, OCA, FRC, PAC, OCC)
1964:114–22). The data were tested for          were included in the principal compo-
outliers using boxplots (not presented).        nent analysis. Only the male sample was
For the non-deformed crania, descriptive        analyzed to prevent sexual dimorphism
statistics were calculated with and with-       from affecting the result, and the female
out outliers. In the deformed sample,           sample was too small to be analyzed in-
the outlying values cannot be regarded          dependently. 24 and 14 individuals could
as random deviations, and the variabil-         be analyzed in the facial and neurocranial
ity should be considered the result of          analyses, respectively.
artificial deformation. Only individual             The deformed crania were analyzed
number 1047, which is believed to have          using Procrustes coordinates to study
a different type of deformation, was ex-        the shape variation across the sample;
cluded prior to the repeated calculation        male and female samples were pooled for
to check whether it was responsible for         this analysis. It included 34 crania with
extreme variability of some traits in the       preserved cranial vaults. Individual num-
male sample.                                    ber 1047 from Baracoa was considered
    Sexual size dimorphism (SSD) for all        an outlier and excluded from the sample
the traits was calculated as the difference     for this analysis for its markedly different
between the mean value of a trait in the        type of deformation.
242                               Taisiya Syutkina et al.

   Additionally, a two-way Mantel test         ed with Asian ancestry. The standard
(Mantel 1967; Sokal and Rohlf 1995)            deviation of most variables in the male
was used to test the association between       sample falls within average standard de-
the matrices of Euclidean craniometric         viations for given morphological traits
and geographical distances for the male        except WFB, XFB, ASB, ALL, ALB, FCD
part of the non-deformed sample. As in         and ALA. In the female sample, these are
previous analyses, morphological vari-         ASB, ZMA, XFB, FCD, ALL and ALB.
ables were divided into two subsets to         However, after identifying and exclud-
avoid a drastic reduction of the sample.       ing outliers (one for each variable), only
The neurocranial subset consisted of six       FCD and alveolar arch dimensions stayed
variables (GOL, XCB, WFB, XFB, AUB,            highly variable within both samples.
ASB) and 20 individuals, while the facial          Despite the female’s sample higher
subset – of seven variables (FMB, ZMB,         variability and lower representativeness,
UFH, OBB, OBH, NLB, NLH) and 23                all the values fall in the same general cat-
individuals.                                   egories (small/medium) as in the male
   All calculations were performed in          sample. The sexual size dimorphism was
R version 4.0.3 (R Core Team 2020)             found to be low among the non-deformed
with RStudio interface version 1.4.1103        crania: only five traits (XFB, PAA, ZMB,
(RStudio Team 2020). The Mantel test           WNB and FCD) proved to have higher
was carried out in R Studio using the          than average rates of SSD in the sample.
function mantel.rtest from package ade4
(Thioulouse et al. 2018), the number of                    Deformed sample
permutations was set at 5000. The geo-         In general, both male and female crania
graphic distances calculations were per-       from the deformed sample (Table 4) have
formed in R using the haversine method         short, very low, and very wide vaults; the
from the package “geodist” (Padgham            frontal bone is flat and wide, it is steeply
and Sumner 2021). The geometric mor-           inclined and often bears a mark left by
phometric analysis was carried out by          the deforming device. Forehead profile
means of geomorph package in R (Ad-            angles values are far beyond the lowest
ams et al. 2021).                              average limits for physiologically normal
                                               crania. All neurocranial and facial widths
                Results                        are very large, so are orbital and facial
                                               dimensions (although orbital and nasal
            Metric variation                   widths tend to be smaller in the female
                                               sample compared to female worldwide
           Summary statistics                  average values). Horizontal profiling of
         Non-deformed sample                   the face is moderate both at nasomalar
Both male and female non-deformed cra-         and zygomaxillary levels.
nia (Table 3) are characterized by medi-          The male sample proved to be very
um-small facial and neurocranial dimen-        heterogeneous: standard deviations of
sions on a worldwide scale (Alekseev           more than a third of variables exceed
and Debets 1964:114–122), including            the world average: XCB, BBH, BPL, XFB,
zygomaxillary and nasomalar facial an-         PAA, OCA, FRC, PAC, OCC, UFH, ALL,
gles, thus suggesting limited expression       FCD, FAN and FAG. Testing for outliers
of cranial metric traits usually associat-     revealed numerous outlying values for
Pre-Columbian Cuba population cranial morphology                    243

the same cranium, 1047, which is be-                   83). Standard deviations decreased after
lieved to be the only one having “tabular              removing it from the sample at this step
erect” deformation type (Rivero de la Cal-             but stayed over the world average for the
le 1949 cited in Herrera Fritot 1964:82–               same variables as before except XFB and

Table 3. Summary statistics for the non-deformed sample
                                                                Male               Female
 Abbreviation                Measurement
                                                           n    Mean    sd     n    Mean    sd
    GOL         Glabello-occipital length                  31   172.7   6.5   14    167.2   5.5
    XCB         Maximum cranial breadth                    27   135.3   3.8   15    130.9   5.1
    BBH         Basion-bregma height                       18   133.9   5.2   10    128.2   4.4
    BNL         Basion-nasion length                       18    97.3   3.8   10     93.5   2.8
    BPL         Basion-prosthion length                    14    93.6   2.8    9     92.1   3.6
    WFB         Minimum frontal breadth                    32    92.8   5.0   14     87.7   4.2
    XFB         Maximum frontal breadth                    28   111.9   5.6   14    108.6   4.8
    AUB         Biauricular breadth                        26   124.0   4.9   14    116.4   4.4
    ASB         Biasterionic breadth                       25   106.0   5.4   13    103.2   4.7
    FRA         Frontal arc                                26   119.0   5.7    9    116.2   4.7
    PAA         Parietal arc                               22   125.3   5.8   10    119.7   4.7
    OCA         Occipital arc                              20   109.9   7.4    6    108.7   2.5
    FRC         Frontal chord                              27   106.7   4.7   10    104.0   2.9
    PAC         Parietal chord                             22   110.1   4.4   10    105.8   4.7
    OCC         Occipital chord                            20    94.5   3.9    6     93.5   2.4
    FMB         Bifrontal breadth                          33   104.3   3.0   13     98.7   3.2
    ZYB         Bizygomatic breadth                        23   132.9   4.2   12    124.5   6.1
    ZMB         Bimaxillary breadth                        28    95.7   3.9   16     90.8   4.8
    UFH         Upper facial height (n-alv)                30    63.1   3.6   14     61.6   1.8
    OBB         Orbital breadth from maxilofrontale        33    41.3   1.8   17     40.0   1.8
    OBH         Orbital height                             34    33.1   1.3   17     33.1   1.5
    NLB         Nasal breadth                              32    23.5   1.7   16     22.9   0.9
    NLH         Nasal height                               32    49.3   3.2   17     47.4   2.3
    ALL         Alveolar length                            14    49.8   3.6    5     49.3   3.9
    ALB         Alveolar breadth                           15    59.2   4.6    5     56.5   3.4
     SIS        Simotic subtense                           31     2.7   0.7   14      2.3   0.4
    WNB         Simotic chord                              31     8.6   1.5   14      7.5   2.0
    MFS         Maxillofrontal subtense                    30     5.7   1.2   14      5.0   1.0
    MFC         Maxillofrontal chord                       30    19.3   2.2   15     17.2   2.0
    FCD         Fossa canina depth                         29     3.8   2.0   15      3.8   2.2
    FAN         Forehead profile angle from nasion         19    82.0   3.8   13     84.5   2.8
    FAG         Forehead profile angle from glabella       19    76.2   3.5   13     78.2   3.2
    GFA         General facial angle                       18    83.1   2.3   13     82.4   2.9
    ALA         Alveolar angle                             14    65.9   6.5    9     65.5   6.0
    NPA         Nasal protrusion angle                     20    21.2   6.6   13     16.9   4.1
    NMA         Nasomalar angle                            29   144.0   4.4   12    144.4   4.1
    ZMA         Zygomaxillary angle                        24   127.2   4.8   13    127.7   6.8
244                                     Taisiya Syutkina et al.

FRC. Female crania seem to be consider-                frontal/facial vertical angles (FAN, FAG,
ably more uniform than the male sample:                GFA, ALA). SSD in the deformed sam-
standard deviations exceed the world av-               ple is not high in terms of metric traits
erage for traits XCB, XFB, PAA, FCD and                (observed index in the sample was high-

Table 4. Summary statistics for the deformed sample
                                                                  Male                Female
 Abbreviation                Measurement
                                                           n      mean     sd     n    mean    sd
      GOL       Glabello-occipital length                  23     172.0    5.6   12    160.6   4.2
      XCB       Maximum cranial breadth                    22     154.5    5.5   12    148.0   5.4
      BBH       Basion-bregma height                       17     125.6    5.4    9    122.3   3.2
      BNL       Basion-nasion length                       17      97.2    4.3    9     91.0   2.1
      BPL       Basion-prosthion length                    16      99.0    5.9    9     92.8   3.1
      WFB       Minimum frontal breadth                    20      96.8    2.6   12     94.2   4.4
      XFB       Maximum frontal breadth                    20     122.5    6.2   10    118.4   5.7
      AUB       Biauricular breadth                        21     136.2    4.2   10    130.0   3.5
      ASB       Biasterionic breadth                       20     112.5    3.4   12    109.1   4.4
      FRA       Frontal arc                                23     117.9    6.1   10    113.0   2.9
      PAA       Parietal arc                               23     107.6    9.7   11    103.4   8.2
      OCA       Occipital arc                              18     113.5   11.7   11    108.5   6.9
      FRC       Frontal chord                              23     110.0    5.4   10    104.8   3.4
      PAC       Parietal chord                             23      95.8    7.0   11     92.5   4.8
      OCC       Occipital chord                            18      97.4    7.6   11     92.5   4.2
      FMB       Bifrontal breadth                          21     109.6    3.3   12    104.8   2.3
      ZYB       Bizygomatic breadth                        17     142.5    4.6   10    132.2   3.7
      ZMB       Bimaxillary breadth                        20     102.1    4.5   11     98.9   2.5
      UFH       Upper facial height (n-alv)                21      70.4    5.1   11     67.4   2.2
      OBB       Orbital breadth from maxilofrontale        22      43.3    1.9   11     41.3   1.8
      OBH       Orbital height                             22      36.1    1.4   11     35.8   0.9
      NLB       Nasal breadth                              22      26.2    1.6   11     25.0   1.6
      NLH       Nasal height                               22      53.8    3.1   11     51.8   1.8
      ALL       Alveolar length                            15      52.4    4.0    7     49.6   2.7
      ALB       Alveolar breadth                           15      64.3    4.1    8     61.2   2.9
       SIS      Simotic subtense                           21       3.6    0.8   11      3.1   0.6
      WNB       Simotic chord                              21      10.4    1.6   11     10.1   1.0
      MFS       Maxillofrontal subtense                    22       6.1    1.2   10      5.8   0.6
      MFC       Maxillofrontal chord                       22      19.4    1.7   10     18.4   2.5
      FCD       Fossa canina depth                         22       3.4    1.4   11      3.1   1.3
      FAN       Forehead profile angle from nasion         16      66.2    5.2   10     67.2   5.9
      FAG       Forehead profile angle from glabella       16      56.4    5.4   10     58.8   5.8
      GFA       General facial angle                       15      81.1    2.9   11     80.5   3.3
      ALA       Alveolar angle                             15      69.8    5.0   11     68.7   6.5
      NPA       Nasal protrusion angle                     16      21.6    4.6   10     18.0   3.0
      NMA       Nasomalar angle                            21     141.2    4.5   11    143.0   3.4
      ZMA       Zygomaxillary angle                        20     127.5    4.2   11    126.5   3.9
Pre-Columbian Cuba population cranial morphology                        245

er than average maximum only for GOL,               plot; moreover, pairs of morphologically
XCB, OCA, OCC and WNB), but male                    similar representatives of both groups are
and female crania are well distinguish-             not infrequent (e.g., individuals number
able visually.                                      18 and 42, 5 and 24, 13 and 25). We can
                                                    also note the central position of the is-
                    PCA                             land individual from Punta del Este, the
          Non-deformed sample                       marginal position of an individual with
Facial variables. The details about the             unknown, but presumably Cuban aborig-
percentage of variance explained and                inal origin, the relative proximity of two
loadings are plotted in Figure 3. As the            crania which come from the same cave in
loading plots indicate, both PCs sum-               Baracoa (individuals number 39 and 14).
marize differences in facial dimensions                 Neurocranial variables. The first two
and nasal height, with nasal breadth and            PCs account for 78% of the total varia-
orbital dimensions being the least sig-             tion of the sample based on neurocranial
nificant. The results of the PCA do not             metric traits (Figure 4). The first PC ex-
reveal any territorial differences in the           plains 49% of the variation and is a size
non-deformed sample. Crania from the                variable as it is negatively correlated with
largest territorial groups – Canímar Aba-           all the variables, length and height of the
jo and Baracoa (which is not a “group” in           vault and occipital arc being the most
a strict sense) are found in all parts of the       significant ones. The second PC explains

Fig. 3. PCA score plot (A) and loadings (B) of seven facial variables. Non-deformed sample. Numeration
    as in Supplement Table 1
246                                     Taisiya Syutkina et al.

Fig. 4. PCA score plot (A) and loadings (B) of ten neurocranial variables. Non-deformed sample. Numera-
    tion as in Supplement Table 1

29% of the variation and is mainly cor-              respectively, both being from the zone of
related with PAA (negatively) and OCA                Baracoa. The PCA based on indices rather
(positively). Although the number of in-             than metric variables yielded very similar
dividuals analyzed has decreased drasti-             results and is not presented here. Indi-
cally due to poorer preservation of crani-           viduals from different geographical loca-
al vault in the sample, it is interesting to         tions appear to be randomly distributed
notice that some trends revealed by the              in these plots, thus not suggesting any
PCA for the facial variables could also be           morphological pattern connected with
observed for the neurocranial data. For              geography in the non-deformed group.
example, individuals from Cueva Fría
(Baracoa) maintain their proximity to                               Mantel test
each other despite being tightly clustered           To further explore whether there is an
with four individuals from four different            association between morphology and
locations. So do individuals number 13               location of the studied crania from the
and 7 also from Maisí (more detailed lo-             non-deformed sample, the Mantel test
cation is unknown), individuals number               was employed over the matrix of Euclid-
16 and 17 from Canímar Abajo. Punta del              ean distances between the morphological
Este individual maintains its central po-            variables and the matrix of geographical
sition, while individuals number 3 and 2             distances between all the sites. No cor-
now hold the most marginal positions in              relation was found between the matrices
terms of the first PC and the second PC,             of neurocranial variables and geograph-
Pre-Columbian Cuba population cranial morphology                           247

ical distances (r = 0.04, p = 0.3). This              the Mantel test did not reveal any con-
set of variables (GOL, XCB, WFB, XFB,                 nection between the craniometric data
AUB, ASB) accounted for the length and                and geographic distances for the non-de-
width dimensions of the crania, but not               formed sample.
their height, due to generally poor pres-
ervation of the bases of the crania. Inclu-                      Deformed sample
sion of the cranial height in the subset                       Shape change analysis
reduced the sample to 12 individuals,                 Figure 5 shows shape variation for the
and although such an analysis cannot                  first two components. Most specimens
be considered reliable, it was carried                in the deformed sample come from the
out to ensure that a possible associa-                same zone of Baracoa, several come
tion was not overlooked. The correlation              from other Cuban locations, five are of
remained negligible and insignificant,                unknown but presumably Cuban origin,
though (r = 0.03, p = 0.3). Finally, the              and three crania come from two sites in
Mantel test on the facial variables yielded           the Dominican Republic. The latter do
a similar result: r = −0.1, p = 0.9. Thus,            not appear to cluster together and are

Fig. 5. Shape variation of the deformed crania in the lateral view. Numeration as in Supplement Table 1
248                                Taisiya Syutkina et al.

scattered across the main cloud of obser-       cantly (p = 0,04) more frequent on the
vations.                                        right side.
    The plot shows a continuous varia-
tion of the deformed crania between the             Discussion and conclusions
shapes at the extremes of the PC1 and
PC2 axes, depicted at the sides of the          The goal of the present paper was to see
plot. PC1, which accounts for 51% of the        whether we could expand our under-
variation, marks the direction of change        standing of the morphological variation
from a more severely affected vault with        of the Cuban pre-Columbian population
flattened frontal and occipital bones and       on the intragroup level by applying var-
a marked depression after the coronal su-       ious methods of studying cranial mor-
ture on the left to a more rounded and          phology.
close to physiological norm vault on the            We acknowledge that the skeletal ma-
right. The crania of both sexes are dis-        terial available for the present study has
tributed randomly across the plot.              not become substantially larger since the
                                                early papers reviewed above. While es-
       Non-metric traits variation              timates of the total number of individu-
Frequencies of the non-metric traits are        als from archaeologically excavated sites
shown in Table 2. Both groups have zero         often numbers into hundreds (Valcárcel
frequencies of the metopic suture, low          Rojas 2016; Rodríguez Hernández 1998;
or zero frequencies of sagittal and cor-        Chinique de Armas and Rodríguez Su-
onal ossicles, ossicles at lambda, Inca         arez 2012), the remains themselves are
bone, palatine tori, pterion types other        usually very fragmentary. The present
than sphenoparietal (although epipteric         study could not cover all existing collec-
bones occur in the 20% of the deformed          tions for various reasons, but much more
individuals), trochlear spine, double con-      representative samples than existing at
dylar facet. Both groups also have high         the moment would be needed for more
rates of the supraorbital notch, multiple       objective conclusions. Relethford (2002)
zygomatic foramina, parietal foramina,          has shown that more than 80% of the to-
posterior condylar canals, and infraor-         tal craniometric variation is found within
bital sutures. At the same time, there is       local populations, i.e., among individ-
a marked contrast in the frequencies of         uals. According to Kozintsev (2016:3),
lambdoid, occipito-mastoidal, squamo-           more than 120 specimens in a sample are
parietal ossicles, and ossicles at the aste-    required to trace its composite nature.
rion between the two groups: all of these       At the same time, if morphological pat-
traits are very common in the deformed          terns are parallelized by some indepen-
group and rather rare in the non-de-            dent criteria, e.g., varying archaeological
formed one.                                     context, the antiquity of the burials etc.,
    No significant differences between          it is justified to suggest a really existing
sexes were found in the frequency of            intragroup differentiation.
non-metric traits in both samples. When             How right are we to pool all non-de-
symmetry was considered, the only sig-          formed crania in one sample? On the
nificant difference was seen in wormian         one hand, our analysis did not reveal
bones in the masto-occipital suture in          any evident clustering of the individu-
the deformed sample: they were signifi-         als in the pooled sample, and we do not
Pre-Columbian Cuba population cranial morphology                           249

possess any certain information about                 from Cave 4 in Punta del Este, a much
them representing different populations.              younger site. Similarly, the absence of
On the other hand, the opposite is also               clear patterns of clustering and scarcity
true: there is no evidence of them be-                of materials does not allow a comparison
ing a single population. Some scholars                between sites with different archaeolog-
(Ulloa Hung and Valcárcel Rojas 2019,                 ical settings. Bolufé Torres (2015) at-
González Herrera 2008) have argued that               tempted to study morphological patterns
“Archaic” or “Siboney” in the Caribbean               among skeletal remains from four sites
is a historical and archaeological con-               situated in the western part of the island
struct, based on first European written               and found significant differences in the
sources and supported by the need for                 orbital breadth between Guayabo Blanco
classification and systematization of ac-             individuals on the one hand and Canímar
cumulated archaeological materials and                Abajo and Cueva Calero – on the oth-
cultures, while in fact this concept masks            er. A geometric morphometric analysis
the actual diversity of groups that form              revealed variation in the cranial shape
this artificial “community”.                          among the three samples. Although the
    In this study, the non-deformed cra-              author concludes that the results might
nia proved to be rather homogeneous,                  indicate the different origin of the in-
with the alveolar region being the most               dividuals, the results should be treated
variable one. Principal component analy-              with caution due to the small sample
sis and Mantel test did not reveal any ter-           sizes (1–8 individuals in each sample).
ritorial differences in the cranial metric            The present study, based on multivariate
traits. No patterns resulting from time               analyses of cranial metrics, does not sup-
differences among the samples could be                port this conclusion.
seen either based on the data available                   The deformed sample is significant-
for the study (see Table 5 for available              ly more variable, especially its male
dates). For the non-deformed sample, in-              part. The principal component analy-
dividuals from the single site of Canímar             sis of the Procrustes coordinates of the
Abajo were found scattered throughout                 cranial vault outline in the lateral norm
the PCA plot, while the individual com-               revealed continuous variability of crani-
ing from Guayabo Blanco, the oldest site              al shapes from the ones with more flat-
among the ones that could be included                 tened frontal and occipital bones to the
in the analysis, was found in the mid-                more curved outlines. In our study, most
dle of the distribution next to the one               of the deformed crania come from the

Table 5. Radiocarbon dates for some of the sites where the materials of the study come from. All dates are
    in calibrated years BP
                Site             Culture           Date                      Reference
Canímar Abajo (Younger cemetery) Archaic        1570–7200                Nägele et al. 2020
Guayabo Blanco                   Archaic        2530–1700                Nägele et al. 2020
Cueva Calero                     Archaic        1500–1100                Nägele et al. 2020
La Caleta (Dominican Republic)   Ceramic        1307–3320               Fernandez et al. 2021
Punta del Este, Cueva 4          Archaic         969–675        Ernesto Tabio, Estudio de las culturas
                                                                          Antiguas en Cuba
El Purial                             Archaic 3644–2780                     Cooper 2007
250                                Taisiya Syutkina et al.

same region of Baracoa (in many cases           suture inside the deformed sample. The
without any additional information), and        only available written evidence of the
three individuals from the Dominican            deforming apparatus in the Caribbean
Republic do not tend to cluster togeth-         (although a long time after the contact
er, so geography cannot account for this        and not directly in Cuba) is provided by
variation of the deformed shapes. More-         Leblond, who evidenced the practice in
over, a highly specific individual number       the late 18th century (1813 cited in van
1047 with the deformation type that has         Duijvenbode 2017:80) and describes a
been described as “tabular erect” instead       procedure of placing two wooden boards
of “tabular oblique” comes from the             at the front and the back of the skull of a
same zone of Baracoa. Sexual differences        baby. The asymmetry found in our study
do not explain the variation either, which      might result from more pressure applied
contradicts the conclusion of Ginzburg          to the right side of the deforming ban-
(1964:202), who reported more mod-              dage when a right-handed person fixes
erate deformation of the female crania.         it.
Thus, it can be assumed that there were             Thus, the study revealed relative ho-
no differences in the application of the        mogeneity of both non-deformed and de-
deforming device to male and female ba-         formed samples and the absence of sys-
bies. The lack of contextual information        tematic differences inside each sample or
and radiocarbon dates for the deformed          proved the impossibility of available cra-
sample make it impossible to test wheth-        niological data to reflect possible intra-
er the variation of the deformed cranial        group differentiation. In other words, by
shapes is due to changes of the tradition       now, there are no grounds for identify-
over time. Our results do not support           ing sub-groups inside the non-deformed
Herrera Fritot’s theory of deforming de-        and deformed samples based on cranial
vices of two types: if this were the case,      morphology.
we would expect to see two separate                 Studies of skeletal remains are inevi-
clusters of crania rather than continuous       tably limited to dealing with samples that
variation. Therefore, the most reason-          do not necessarily reflect the properties
able explanation for the observed trend         of the population it represents, but the
is an individual variation of the resulting     more numerous the sample is and the
shapes.                                         more information about it is available,
    Variation of non-metric traits in each      the more credible the conclusions can
sample found in this study is totally in        be. It is probable that as the pool of ma-
accordance with earlier works (Rivero           terials increases, the trends identified in
de la Calle 1983:180–181). The high             the present study will be proved errone-
frequency of sutural ossicles in the de-        ous, and new patterns not evident in the
formed sample can be both due to the            analysis of such limited material will be
effect of deformation or manifesta-             revealed instead. We hope that this paper
tion of a particular genetic complex – a        will serve as a reference point for future
non-deformed sample of the same or-             studies that will confirm or refute the
igin would be required to make a defi-          preliminary conclusions offered here and
nite conclusion. An interesting result is       continue to explore relations between
an asymmetry seen in the expression of          different groups populating the island
wormian bones in the mastoid-occipital          throughout its pre-Columbian history.
Pre-Columbian Cuba population cranial morphology                   251

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        Corresponding author
                                                  craneofaciales en las poblaciones pesca-
                                                  doras recolectoras cazadoras del occiden-
Taisiya Syutkina, Checherskiy proezd-             te de Cuba: Cueva del Perico I, Canímar
24-­130, Moscow, Russian Federation               Abajo, Cueva Calero y Guayabo Blanco.
e-mail: syuttaya@gmail.com                        Tesis en opción al grado de Licenciado en
                                                  Biología. La Habana: Universidad de La
                                                  Habana.
252                                    Taisiya Syutkina et al.

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