Journal of Archaeological Science xxx (2018) 1e10

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Journal of Archaeological Science

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Archaeology, biology, and borrowing: A critical examination of

Geometric Morphometrics in Archaeology
Mercedes Okumura a, *, Astolfo G.M. Araujo b, c
a
Laborato
rio de Estudos Evolutivos Humanos, Departamento de Gen
etica e Biologia Evolutiva, Instituto de Bioci^ ~o Paulo, Brazil
encias, Universidade de Sa
b ~o Paulo, Brazil
Instituto de Estudos Avançados, Universidade de Sa
c
Museu de Arqueologia e Etnologia, Universidade de Sa~o Paulo, Brazil

a r t i c l e i n f o a b s t r a c t

Article history: Geometric Morphometrics (GM) is a method originally applied in Evolutionary Biology studies, using the
Received 17 June 2016 analysis of change in size and shape in order to better understand ontogenetic sequences, phylogenetic
Received in revised form relations, among other issues. The application of GM in archaeological materials has seen a sharp in-
13 September 2017
crease in the last decade, mostly associated with theoretical approaches from Evolutionary Archaeology.
Accepted 24 September 2017
Available online xxx
This is not an isolated case, since most methods used by Evolutionary Archaeologists have been bor-
rowed from Biology, provoking discussion with regard to the future development of Evolutionary
Archaeology and its methods (Lycett, 2015). This article aims to discuss some concepts that have been
Keywords:
Geometric Morphometrics
directly borrowed from the application of GM in Biological Sciences and that have not been subject to
Evolutionary Archaeology much thought when used in Archaeology. Such concepts include homology and landmark types, the
Stone tools concept of modularity, as well as the idea of allometry. As much as archaeologists using GM can learn
Interdisciplinarity from past discussions held by biologists regarding the above mentioned concepts, it is high time for
archaeologists to further discuss ideas concerning the use of these concepts in archaeological studies.
© 2017 Published by Elsevier Ltd.

1. Introduction Ecology, including Optimal Foraging Theory, widely applied by ar-

1.1. Borrowing across disciplines: from biology to the humanities to
Archaeology
“I did not say that the borrowing of concepts, tools, methods, or
models from other sciences is necessarily bad. What is at issue is
the appropriateness of the model for the job at hand.” (Service,
1969: 77).
The borrowing of concepts, tools, methods, or models from
Biology to the Humanities and ultimately to Archaeology is not a
new enterprise. The very concept of (natural) selection by Darwin is
a borrowing from the activities set forth by animal breeders to the
biological realm (Evans, 1984), developed and transformed by bi-
ologists to fit their disciplinary purposes, and then returning to the
human sphere due to the realization that the same general ratio-
nale can be applied to human affairs (Dunnell, 1980).
Probably the most popular instance of borrowing from Biology
to Archaeology would be the models originated in Evolutionary
* Corresponding author.
E-mail address: okumura@ib.usp.br (M. Okumura).
chaeologists to the study of prehistoric hunter-gatherer groups
(Bettinger and Baumhoff, 1982; Broughton, 2002; Byers and Ugan,
2005; Raab, 1992; Smith and Winterhalder, 1981, among others).
The discussion about dangers of borrowing concepts, tools,
methods, or models from other disciplines is also not a new one,
although it might have received less attention than it ought to have.
Back in 1969, Service (1969) alerted to the dangers of mouthtalk
(defined by the author as the mindless borrowing of concepts) as
well as the danger of borrowing techniques without concerns about
the problem to be tackled. According to Keene (1983), the problem
begins when there is the adoption of new techniques without re-
gard for their original intent or limitations. In that way, similarities
between domains are reinforced and differences are either mini-
mized or disregarded, leading to a lack of adaptation of such
techniques to the new discipline, even when that is a necessary
step. The author stresses that usually the lending discipline will set
the research agenda, carrying (mostly in an implicit way) a his-
torical legacy that includes its own particular worldview. The rea-
sons for adopting concepts, tools, methods, or models from other
disciplines are usually related to the perceptions that the lending
science can produce more sophisticated and accurate analytical

https://doi.org/10.1016/j.jas.2017.09.015
0305-4403/© 2017 Published by Elsevier Ltd.

Please cite this article in press as: Okumura, M., Araujo, A.G.M., Archaeology, biology, and borrowing: A critical examination of Geometric
Morphometrics in Archaeology, Journal of Archaeological Science (2018), https://doi.org/10.1016/j.jas.2017.09.015
2 M. Okumura, A.G.M. Araujo / Journal of Archaeological Science xxx (2018) 1e10
tools than the borrowing science (Keene, 1983). These foreigner
items can also give a more scientific aura to the research (Service,
1969), something pursued by some theoretical approaches in
Archaeology (Binford, 1962, 1964, 1965, 1987, Watson et al., 1971,
1984; but see Dunnell, 1982, 1992; Hunt et al., 2001; Lyman et al.,
1997, for a critique). Therefore, in some cases, such items are first
borrowed and then a question has to be made in order to apply
them (Service, 1969), which is the opposite of what should be done
(Sherif and Sherif, 2009: 12). On the other hand, borrowing across
disciplines is also a pioneer activity, paying its price accordingly. A
rather small number of scholars are prone to “jump over the fence”
of disciplinary boundaries and pursue useful scientific rationales
outside their “grass field”, and such ideas are often dismissed at
face value by colleagues due to lack of proper knowledge or interest
in the subject matter. This is even more critical in the Humanities,
where the fear of the “random, chance, and unguided” (e.g.,
Sandstrom, 2013) or simply a perceived danger of losing control
over discipline boundaries (e.g., Halloway, 1969; see Wallerstein,
2003) are the most commonly raised issues. Interestingly, bi-
ologists borrowing human concepts such as “intentions”, “desires”
and “beliefs” to explain cell biochemistry do not show this fear
(Jonker et al., 2002).
In the last decade, Geometric Morphometrics (GM) e a method
developed mainly by evolutionary biologists e has seen a sharp
increase in its application to archaeological questions. As any other
method borrowed from Biological Sciences, some of its concepts
might need to be adapted for its use in archaeological materials.
Which GM concepts were or should have been addressed by ar-
chaeologists, how the adaptation of such concepts was or was not
made, as well as the subsequent implications of such a process are
the main subject of this article.
1.2. A general overview of GM application in material culture
GM has been applied to several instances of archaeological
materials in order to better understand and describe shape varia-
tion. Obviously, due to the common questions shared with Evolu-
tionary Biology, the greater amount of case studies involving the
use of GM will refer to research involving biological affinities in
human archaeological remains. Cranial morphology has been
widely used to reconstruct evolutionary relationships in archaeo-
logical groups (de Azevedo et al., 2011, 2017; Galland, 2015; Galland
and Friess, 2016; Gonza
lez-Jose
et al., 2003, 2008; Manríquez et al.,
2011; Martínez-Abadías et al., 2006; Neves et al., 2005; Perez et al.,
2009; Pinhasi and von Cramon-Taubadel, 2009; von Cramon-
Taubadel and Pinhasi, 2011). Besides this, GM has also been used
in human remains from archaeological contexts to investigate
variation in artificially deformed crania (Friess and Baylac, 2003;
Perez, 2007). Other uses of GM that are relevant for archaeolo-
gists working with human remains are those related to sex and age
estimation (Franklin et al., 2007, 2008; Gonzalez et al., 2009, 2011,
among others). The focus of this review will be the application of
GM in archaeological artefacts (material culture) and how archae-
ologists working with material culture have been dealing with this
borrowing.
The term “material culture” is part of the bedrock in the disci-
pline of Archaeology and considered the uniting feature combining
and relating all the different practices together (Oestigaard, 2004).
Therefore, it comes as no surprise that some archaeologists,
knowing the explanatory potential of GM, have been applying it to
material culture in order to better understand their variation in
time and/or space (Archer and Braun, 2010; Buchanan and Collard,
2010a; Buchanan et al., 2014; Cardillo, 2006, 2009, 2010; Cardillo
~ eira et al., 2011, 2012; Charlin and Gonza
et al., 2016; Castin
lez-

, 2012; Charlin et al., 2014; Costa, 2010; Davis et al., 2015; de
Jose
Please cite this article in press as: Okumura, M., Araujo, A.G.M., Archaeology, biology, and borrowing: A critical examination of Geometric
Morphometrics in Archaeology, Journal of Archaeological Science (2018), https://doi.org/10.1016/j.jas.2017.09.015
Azevedo et al., 2014; Franco et al., 2009; Iovita, 2011; Iovita and
McPherron, 2011; Lycett and Von Cramon-Taubadel, 2012; Lycett
et al., 2006, 2010; Okumura and Araujo, 2013, 2014, 2016; Picin
et al., 2014; Selden et al., 2014; Serwatka, 2015a; Shott and Trail,
2010; Thulman, 2012; Wang et al., 2012). Although the use of GM
applied to material culture does not necessarily involve a theoret-
ical connection to Evolutionary Archaeology, most case studies
show this relation, which obviously has to do with the common
interests shared by Evolutionary Biologists and Evolutionary Ar-
chaeologists, including essential concepts like selection, drift, and
transmission, as well as the idea that both Evolutionary Biology and
Archaeology are historical sciences (Dunnell, 1980, 1982; Eerkens
and Lipo, 2007; Neiman, 1995; O'Brien et al., 2010; Shennan,
2000, 2001). The borrowing of theoretical concepts and methods
from Evolutionary Biology by Evolutionary Archaeologists is not
restricted to GM. Other methods, created by and widely applied in
Evolutionary Biology, such as phylogenetic methods (mainly Cla-
distics), have also been borrowed by Evolutionary Archaeologists
(Beck and Jones, 2007; Buchanan and Collard, 2007, 2008a, 2008b;
Collard, 2010; Collard and Shennan, 2000; Collard et al., 2006;
Coward et al., 2008; Darwent and O'Brien, 2006; Eerkens et al.,
2006; Harmon et al., 2006; Jordan and Shennan, 2003; Lycett,
2007, 2009a, 2009b; Marwick, 2012; O'Brien and Lyman, 2003;
O'Brien et al., 2001, 2002, 2012; Prentiss et al., 2011; Rivero, 2016;
Rivero and O'Brien, 2014; Rorabaugh, 2012; Tehrani and Collard,
2002, 2009; Temkin and Eldredge, 2007; Tolstoy, 2008). This sce-
nario has caused some concern with regard to the future devel-
opment of Evolutionary Archaeology and its methods (Lycett, 2015,
see also O'Brien et al., 2002 for a thoughtful discussion on some
important issues regarding the use of cladistics in archaeological
materials).
2. GM key concepts: from biology to Archaeology
Like any other method, there are concepts which are funda-
mental in applying GM to a given set of structures. These ideas,
which are ultimately derived from the Biological Sciences, are
related to theoretical expectations about homologous structures,
morphological integration, as well as allometric relations. Ideally,
the use of GM applied to archaeological artefacts should be pre-
ceded by a thoughtful discussion by archaeologists about the im-
plications of borrowing such concepts, if they are adequate for use
in archaeological studies and if not, which would be the feasible
alternatives to them. In this section, we will discuss how land-
marks, modularity, and allometry have (or have not) been
addressed by archaeologists when working with GM applied to
material culture.
2.1. Landmark types and the concept of homology in GM applied to
Archaeology
Geometric morphometrics can be defined as the quantitative
representation and analysis of morphological shape using geo-
metric coordinates. Therefore, morphometricians are interested in
data that captured the geometry of a given morphology, as well as
in developing methods to analyze such data. This included methods
for both landmark and outline data (Adams et al., 2004).
Landmarks are “samples of discrete points which correspond
among all the forms of a data set” (Bookstein, 1990: 63). According
to Zelditch et al. (2004: 24), landmarks should not change their
topological positions relative to other landmarks (a one to one
correspondence in the specimens to be compared, Viscosi and
Cardini, 2011), should result in a good coverage of the studied
morphology, be observed repeatedly and reliably, and present
coplanarity. Although in recent years new approaches have been
 M. Okumura, A.G.M. Araujo / Journal of Archaeological Science xxx (2018) 1e10 3

created to overcome drawbacks regarding landmarks in GM (Aneja limitations, geometrical homology is based on the correspondence
et al., 2015; Pomidor et al., 2016), most GM studies in Archaeology of positions across specimens, partially inspired by D'Arcy
still use them. The early system of landmark classification Thompson (1942) grid transformations (Jardine, 1969). In a way,
comprised three types of landmarks (Bookstein, 1991: 63e65) and geometrical homology and biological homology share the basic
it was the subject of much discussion in the early days of GM given empirical criterion of correspondence in relative position proposed
the problems related to different homology types (biological, geo- in some traditional definitions of biological homology (Darwin,
metric, etc) and to the identification of potential directions of forces 1859: 434e5; Owen, 1848: 175e6). This concept was adopted,
acting in a given structure (O'Higgins, 2000; Slice, 2005; Zelditch relabelled as “topographic homology”, and further developed much
et al., 2004: 31).; later by Jardine (1969), who claimed that using only correspon-
An alternative to the use of landmarks are outlines. There are dence in relative position was not a satisfactory criterion.
three basic ways of analysing outlines: Fourier analysis, eigenshape, The discussion about whether or not morphometric variables
and semi-landmarks (Lawing and Polly, 2010). Fourier analysis uses can express biological homology was further addressed by
sine and cosine harmonic functions to describe the positions of MacLeod (2002). There are two main questions pointed out by the
outline coordinates (Rohlf, 1986). In eigenshape, the coordinate author and they refer, respectively, to structure-level homology and
points of a given outline are converted to a phi function, which is a point-level homology. The first question is whether a given set of
list of the angles from one point to the next one in the series landmark points based on biological elements is homologous to
(MacLeod, 1999, 2008). Finally, outlines can be analysed using another set of landmark points defined in a similar way. The second
semi-landmarks (also named sliding landmarks), which are the is whether individual landmark points are homologous with other
points located relative to one another by some consistent rule (e.g., landmark points. Because of the traditional landmark definition
equal linear spacing from preceding point, equi-angular spacing used in GM, corresponding landmarks are, by definition, geomet-
according to a radius vector originating from the centroid of a rical homologues. The author recalls that in most cases many
closed form), that collectively trace the geometry of a form different sets of landmarks will pass the similarity, conjunction, and
(Bookstein, 1991, 1997). The concept of semi-landmark was pro- congruence tests of biological homologues, rendering it impossible
posed by Bookstein (1997) as a way to circumvent the problem of to distinguish true homology (there can only be one pair of land-
biological homology observed in the “classical” landmark system. mark points that define a length on any biological structure that are
Although many authors have shown concern about the lack of biological homologues of another pair of landmark points on
homology of each point in outline methods (Bookstein et al., 1982; another homologous structure) from false homology. Although
Gunz et al., 2005), some authors consider it a minor problem, Type 1 landmarks (sensu Bookstein, 1991) may show a coincidence
because the total outline structure will be either biologically ho- between biological and topological homology (Wagner, 1994), they
mologous or functionally analogous and the greatest importance represent a minority of landmark types used for morphometrical
should be placed on the overall structure shape, not on individual analyses. For MacLeod (2002), the lack of proper developmental or
points (Ehrlich et al., 1983; Lawing and Polly, 2010; MacLeod, 1999, phylogenetic evidence makes the point homologies in GM a non-
2008; Read and Lestrel, 1986; Zelditch et al., 1995). The conceptual testable assumption.
problems regarding the lack of biological homology in some land- Most GM studies in Archaeology have used a combination of
mark types, as well as in semi-landmarks, was overcome by an landmarks and semi-landmarks or only landmarks (Fig. 1,
alternative definition given by MacLeod (1999, 2008). For the Buchanan, 2006; Buchanan et al., 2007; Buchanan and Collard,
author, landmarks are coordinate locations that bear some topo-
logical, anatomical, functional, etc. Correspondence across all
structures and it makes no sense to attribute homology to specific
points, because homology would be a property observed in entire
structures. The author also considered as landmarks evenly spaced
points whose placement is arbitrary with respect to anatomical
traits. Therefore, for MacLeod, landmarks would include all Book-
stein's landmark types, as well as semi-landmarks. It might be that
the main problem regarding the presence or absence of homology
in landmark types refers to a clear definition of that term (Lycett,
2009c) and probably there are at least two different (albeit
related) concepts of homology being used by different researchers
(Gunz et al., 2005). The classic biological definition of homology
(considered the hierarchical basis of comparative biology, Hall,
1994: 1), refers to morphological structures in biological beings
that present similarities of structure, physiology or development
related to a shared evolutionary ancestry, meaning that only
explicit entities of selection or development can be considered
homologous (Gunz et al., 2005). Although such a traditional defi-
nition seems to be able to encompass many different biological
areas, in reality a single definition of homology cannot be applied to
all elements and all levels in the hierarchy of biological organiza-
tion (which includes molecules, genes, cells, organs, embryos, or-
ganisms, populations, communities, etc, Hall, 1994: 1). Leaving
aside that discussion, there is what some authors call “geometric
homology”, which is a concept related to GM, where homology
becomes synonymous with morphological points presenting cor-
respondence, clearly ruling out any possibility of such points being Fig. 1. Example of use of landmarks (large dots) and semilandmarks (small dots) in
entities of selection (Gunz et al., 2005). Regardless of such stemmed bifacial points (adapted from Gonza
lez-Jose

and Charlin, 2012).

Please cite this article in press as: Okumura, M., Araujo, A.G.M., Archaeology, biology, and borrowing: A critical examination of Geometric
Morphometrics in Archaeology, Journal of Archaeological Science (2018), https://doi.org/10.1016/j.jas.2017.09.015
4 M. Okumura, A.G.M. Araujo / Journal of Archaeological Science xxx (2018) 1e10
Fig. 2. Example of homology among landmarks in different point types (adapted from Okumura and Araujo, 2014).
2007; Cardillo, 2010; Charlin et al., 2014; Costa, 2010; Lenardi and
Merwin, 2010; Okumura and Araujo, 2013, 2014, 2016; Scartascini
et al., 2009; Shott and Trail, 2010; Thulman, 2012), and a few
have used Fourier analysis (Cardillo, 2010; Fox, 2015; Friess and
Baylac, 2003; Gero and Mazzullo, 1984; Iovita, 2009, 2010;
Serwatka, 2015b). Although homology is one of the basic con-
cepts to define the optimality of landmarks, there has been very
little discussion regarding the implications of landmark and semi-
landmark uses in GM studies applied to archaeological material
culture. One exception to this point would be the works by Lycett
(2009c; Lycett et al., 2006; Lycett and Chauhan, 2010; Lycett and
von Cramon-Taubadel, 2013), one of the pioneers to discuss prob-
lems related to applying GM methods to artefacts, recalls that most
lithic artefacts do not present many obvious identifiable “homolo-
gous” (correspondent) points. For the author, overcoming this
obstacle and demonstrating the potential of applying GM to answer
archaeological questions was the main challenge facing the science
of lithic morphometric analysis. The author also proposes using
cladistics to test the hypothesis of morphological homology (cor-
respondence of form) being equivalent to phylogenetic homology
(due to descent via a common ancestor, Lycett, 2007) in lithic ar-
tefacts. However, in most of the GM studies applied to Archaeology,
the proposed homologies are either not given much attention or
always considered true, and therefore, not feasible to be tested
(Barrientos, 2010). The consequence is that homology on stone
artefacts is essentially based on the similarity criterion (Barrientos,
2010).
Likewise, Shott and Trail (2010) also raised awareness as to the
implications of landmark types for Archaeology. In order to avoid
the obvious problems of using Bookstein's landmark classification
in archaeological materials, the authors include the landmark
classification proposed by MacLeod (1999, 2008). Such approach
can also be found in the works of Lycett and colleagues (Lycett and
Chauhan, 2010; Lycett and von Cramon-Taubadel, 2013). Shott and
Trail (2010) use lithic projectile points to discuss the utility of
landmark definitions. Projectile point tips and base corners, for
example, can be considered discrete and functionally meaningful
points, presenting geometric homology (Fig. 2). For the authors, the
use of a few landmarks combined with many semi-landmarks in
stone tools that lack distinct haft and blade elements (such as
Acheulean bifaces) is also problematic if archaeologists take into
account discussions regarding semi-landmark conceptual prob-
lems (but see Mitteroecker and Gunz, 2009; among other authors,
for a different view). Given such problems, Cardillo (2010) defends
the study of morphological structures regardless of information on
their homology and that such homology hypothesis can be
addressed through the application of phylogenetic methods. For
the author, archaeological artefacts cannot possibly present bio-
logical homologies, being the result of standardized cultural
Please cite this article in press as: Okumura, M., Araujo, A.G.M., Archaeology, biology, and borrowing: A critical examination of Geometric
Morphometrics in Archaeology, Journal of Archaeological Science (2018), https://doi.org/10.1016/j.jas.2017.09.015
practices, whose creation, transmission and extinction can be
explained by Cultural Transmission Theory (Cavalli-Sforza and
Feldman, 1981; Boyd and Richerson, 1985). This view is also
shared by Barrientos (2010), who presents the concept of homology
as a correspondence of points across lithic artefacts, not necessarily
implying a common evolutionary or developmental pathway. The
author justifies his view, recalling that one of the aims of GM
studies is to create meaningful information on morphological
variation across specimens (O'Higgins, 2000), regardless of their
genealogical relationships. Actually, the way landmark or outline
sets from a biological element relate to real homologous structures
is quite difficult to access due to the frequent combination different
types of landmarks (sensu Bookstein, 1991, 1997) and therefore
different “levels” of homology (Barrientos, 2010). Cardillo (2010)
proposes that in the case of archaeological artefacts, the choice of
points must be related to research questions, as well as to the to-
pological particularities of artefact types, and technical and
morphological criteria. A key issue here is that as Lycett has
repeatedly pointed out (Lycett, 2009c; Lycett and Chauhan, 2010),
there are several ways of defining “homology” both in biology and
in archaeology (geometrically, developmentally, and phylogeneti-
cally), and as such, it is important to be clear about how the term is
being used in a given case both analytically and theoretically.
2.2. Modularity and morphological integration in archaeological
artefacts
Sizes and shapes of different parts of organisms are coordinated
to make up a functional whole. The concept of integration and
modularity was of pivotal importance to the beginnings of the
study of morphology and development by pioneers like Georges
Cuvier1 and Wilhelm Roux in the 19th century (Mayr, 1959; Raff
and Wolpert, 1996). The present concepts of morphological inte-
gration and modularity were created in the middle of the 20th
century (Olson and Miller, 1999) and integrated with ideas from
other areas, such as Evolutionary Genetics and Developmental
Biology. Analyses of morphological integration and modularity
have been conducted with GM approaches for over a decade and
both have become central concepts in Evolutionary Biology as well
as GM (Klingenberg, 2013). Morphological integration and modu-
larity are often presented as complementary concepts in the liter-
ature and groups of correlated variables are usually interpreted as
modules. However, in Evolutionary Biology, morphological inte-
gration refers to the decomposition of morphological traits
1
Although Cuvier considered the organisms parts as so tightly integrated
(indecomposable wholes) that he basically assumed a lack of modularity in most
cases (Schlosser and Wagner, 2004).
 M. Okumura, A.G.M. Araujo / Journal of Archaeological Science xxx (2018) 1e10
according to their phenotypic covariance structure (meaning, the
covariation among morphological traits, Klingenberg, 2013). Usu-
ally such decomposition is compared across adult and/or subadult
individuals belonging to a single taxon or across many taxa
(Mitteroecker and Bookstein, 2007) and is interpreted in terms of
developmental and evolutionary constraints (Cheverud, 1984;
Futuyma, 2010; Klingenberg, 2008, 2013). The interactions that
constitute the mechanisms responsible for morphological integra-
tion cannot usually be directly observable, but the underlying
mechanisms can be inferred from the covariation of morphometric
measurements and hypotheses about their effects can be tested
(Klingenberg, 2013). Modularity is a property of complex structures
or processes that are either experimentally or conceptually sepa-
rable into several “near-decomposable” modules (Simon, 1962).
Modules are complexes of parts that are closely integrated inter-
nally, but are relatively independent of other complexes
(Klingenberg, 2008, 2013). Such modules usually reflect develop-
mental, functional, or evolutionary processes. Therefore, modu-
larity can be characterized as a property of processes, while
morphological integration is a particular pattern of variation and
covariation among parts that these processes brought about
(Mitteroecker and Bookstein, 2007). In any case, modularity is a
concept that is closely related to morphological integration
(Klingenberg, 2008).
GM offers a variety of methods to address a wide range of hy-
potheses concerning modularity (Klingenberg, 2013), which can be
related to anatomical, developmental, functional or genetic ques-
tions. Such hypotheses are defined in terms of the landmarks that
belong to the hypothesized modules. The relative positions of
landmarks from the same module should be relatively constant,
showing a high integration. On the other hand, if the landmarks are
divided into subsets that do not coincide with the putative mod-
ules, the high covariation observed within each module contributes
to the covariation among subsets of landmarks, making the general
covariation among subsets stronger (Klingenberg, 2013). This
pattern of covariation can be tested in empirical terms, through the
quantification of covariation among the groups of landmarks from
different postulated modules and the comparison between such
covariation and the covariation among subsets of landmarks that
have been divided in different ways (Klingenberg, 2009).
The idea of modularity has always played an important role in
disciplines like art, engineering, architecture, computer science and
social sciences including evolutionary psychology (Cosmides and
Tooby, 1992). For each discipline, specific operational criteria of
what is a module have been (or should have been) devised
(Mitteroecker and Bookstein, 2007). In Archaeology, the concept of
modularity in GM has not been extensively explored. Cardillo
(2010) discusses very briefly the possibility of dividing the projec-
tile point morphology into a set of modules based on morphological
or technological criteria. In this case, correlation patterns between
modules (for example, between the blade and the neck or base of a
projectile point) could be related to functional integration of
different sections of artefacts. Very few authors have further
developed the concept of modularity in GM studies applied to
material culture (Charlin and Gonza
lez-Jose

, 2012; de Azevedo
et al., 2014; Gonza
lez-Jose

and Charlin, 2012). These authors use
the borrowed concept of modularity in GM from Biological Sciences
and discuss how that can be applied to further understand function
in prehistoric lithic points from southern Patagonia used by
different systems (arrows, thrown spears, and hand-held points),
when points are considered to present two modules (blade and
stem). Modules were defined in terms of internal relations/
dependent elements (like size and shape characteristics of the stem
or blade), as well as external relations/independent elements (the
blade in relation to the rest of the point); such definitions are
Please cite this article in press as: Okumura, M., Araujo, A.G.M., Archaeology, biology, and borrowing: A critical examination of Geometric
Morphometrics in Archaeology, Journal of Archaeological Science (2018), https://doi.org/10.1016/j.jas.2017.09.015
5
clearly borrowed from the concept of modularity in Biological
Sciences (Schlosser and Wagner, 2004). However, the authors
(Gonz

and Charlin, 2012) propose a more specific inter-
alez-Jose
pretation of modularity applied to material culture (in this case, to
stone tools), stating that modularization might be economically
advantageous because it simplifies the design, the making, and the
maintenance of stone tools (Bleed, 1986; Krohs, 2009). In fact, the
modular design is one of the key characteristics of maintainable
systems, according to Bleed (1986), when discussing the design of
hunting weapons. The way modular systems can evolve in artefacts
is presented using the same hypotheses of modularity in biological
systems: by the division of a highly integrated system or by the
integration of different systems (Wagner and Altenberg, 1996). The
authors propose that modularity in most non composite lithic
points are cases of specialization of previously established struc-
tures (the initial core). They test the hypothesis that such points can
be considered as a two-module system formed by the blade and the
stem (usually considered as techno-functional units), and they aim
to evaluate the degree in which shape, size, asymmetry, blade:stem
length ratio, and tip angle might explain the observed variance and
differentiation among points supposedly aimed to accomplish
different functions. Results point out that, in the analysed sample,
the blade and the stem can be considered as modular structures
when the effects of reduction are included. However, resharpened
points tend to present a modular pattern in terms of tip/rest of the
point. Another interesting result is that the three studied types
(arrows, thrown spears, and hand-held tools) show different
modular organization. In other words, shape, size, asymmetry,
blade:stem length ratio, and tip angle can be important elements to
infer the function of these lithic artefacts (Gonz

and
alez-Jose
Charlin, 2012). The implications of such findings are that the
modular behaviour observed on the blade and on the stem would
be expected when functional demands are greater than design
demands. Another important implication is that a modular
behaviour would greatly decrease the costs of experimenting with
new designs, making it possible to modify parts of a system without
changing it completely.
2.3. Allometry as an integrating morphological factor
In GM, the form of an object is described as size plus shape.
These two elements are defined relative to each other: shape is
defined as all the geometric information that remains when loca-
tion, scale and rotational effects are filtered out from an object
(Bookstein, 1998). Size is often equivalent to “centroid size” which
is the square root of the sum of the squared distances from all the
landmarks to the centre of the form (the centroid). Centroid size is a
measure of size that is mathematically independent of shape
(Zelditch et al., 2004: 13), and it is not equivalent to any distance
between landmarks (Bookstein, 1991: 95). Most importantly, size
and shape can be separated in mathematical terms (Darroch and
Mosimann, 1985) and its relationship can be further explored
(Klingenberg, 2013). Allometry is defined as the relationship be-
tween size and shape and it can be a key factor to understand
morphological integration (Klingenberg, 2009; Klingenberg et al.,
2001; Mitteroecker and Bookstein, 2007). There are different
levels of allometry: static, ontogenetic, and evolutionary allometry
(Cheverud, 1982; Cock, 1966; Gould, 1966; Klingenberg and
Zimmermann, 1992; Klingenberg, 1996, 1998) and such levels are
defined by the process that produces the size variation involved in
the allometric effects: individual variation within populations at a
given ontogenetic stage for static allometry, growth for ontogenetic
allometry, evolutionary change of size for evolutionary allometry
(Klingenberg, 2013).
Given that the relationship between size and shape is often
6 M. Okumura, A.G.M. Araujo / Journal of Archaeological Science xxx (2018) 1e10
linear or nearly linear, allometry results in shape variation that is
mainly directed towards one direction in shape space. The combi-
nation between great size variation and strong allometry, will
imply that most of shape variation will be composed by such size-
related component of shape variation. Strong allometry (when
compared to other sources of variation) will result in a great inte-
gration of shape variation, potentially affecting the analysis of
integration and modularity (Klingenberg, 2009, 2013). Because
allometry can affect many parts of a structure or organism together,
it will result in a combined change of the whole configuration of
given landmarks (Klingenberg, 2013).
In GM applied to the same type of archaeological artefact, usu-
ally the greater source of variation among individuals will be a
consequence of size differences (Lycett, 2009c). The removal of
artefact size is a usual technique among GM studies in Archaeology,
which ensures that differences among artefacts will be based on
shape characteristics rather than general size. That is particularly
important when dealing with stone tools, because the size of a
given raw material (blank form) will limit the size of the finalized
artefact (Chauhan, 2003). According to Lycett (2009c), size removal
allows GM analysis to focus on comparison among artefact shapes.
However, the author stresses that such a removal does not signify
size being treated as irrelevant in order to understand variation
among stone tool morphology, but that it should be analysed and
clearly defined apart from shape. In fact, it is not uncommon to see
the correlation between vectors of size and shape of artefacts being
referred as allometry by some archaeologists (Andrews et al., 2015;
Archer and Braun, 2010; Buchanan, 2006; Okumura and Araujo,
2014).
Another very promising theme involving allometry and stone
tools is the role of lithic resharpening or rejuvenation practices that
prolong the use of lithic artefacts (Dibble, 1984; Shott and Ballenger,
2007). Such practices are closely related to the production, use,
maintenance and discard (Schiffer, 1972; Shott, 2005) of stone tools.
Resharpening can be defined as a maintenance process, usually
observed when the finalized artefact was extensively used,
implying that it was blunt or damaged (Kooyman, 2000:2).
Resharpening in stone tools usually aims to produce a fresh, sharp
cutting edge in a dulled tool (Hayden, 1987). Some authors use the
term rejuvenation as the refurbishing of a broken tool into a
functionally equivalent tool (Towner and Warburton, 1990).
Therefore, resharpening and rejuvenation of stone tools can
potentially modify their dimensions, including length, width and
thickness (Hoffman, 1985). Such a reduction in size related to such
practices lead some authors to propose the concept of stone tool
ageing or senescence (Lycett, 2010; Shott, 2010). Different types of
stone tools, including scrapers (Andrews et al., 2015; Dibble, 1984),
handaxes (Archer and Braun, 2010; Iovita and McPherron, 2011;
McPherron and Dibble, 1999), and projectile points (Buchanan,
2006; Buchanan and Collard, 2010b; Charlin and Gonz

,
alez-Jose
2012; de Azevedo et al., 2014; Flenniken and Raymond, 1986;
Gonz
alez-Jose
and Charlin, 2012; Okumura and Araujo, 2014;
Shott et al., 2007) can be modified through resharpening and
rejuvenation practices. In the case of lithic stemmed projectile
points, the blade will be the region that presents the greater po-
tential to be modified by resharpening or rejuvenation in com-
parison to the stem (Fig. 3, Charlin and Gonza
lez-Jose
, 2012).
Similarly, in non-stemmed projectile points, bases are rarely
resharpened (Ahler and Geib, 2000; Musil, 1988). For example, for
some authors, the great variation observed in North American
Clovis points might be related to resharpening or rejuvenation
practices (Hofman and Graham, 1998; Titmus and Woods, 1991, but
see Buchanan et al., 2014, 2015 for a different view). Given the
reductive nature of such processes, decrease in size is a by-product
that can result in strong allometry (Cardillo, 2010). Allometry is
Please cite this article in press as: Okumura, M., Araujo, A.G.M., Archaeology, biology, and borrowing: A critical examination of Geometric
Morphometrics in Archaeology, Journal of Archaeological Science (2018), https://doi.org/10.1016/j.jas.2017.09.015
Fig. 3. Example of allometric changes related to resharpening in GM analysis applied
to stemmed bifacial points from southern Brazil.
more widely discussed in relation to raw material availability and
reduction intensity (including resharpening practices), but other
elements including function, cultural tradition, as well as cognitive
and/or biomechanical differences can also be potentially important
factors affecting form variability (Lycett, 2009c) and therefore,
allometry.
3. Conclusions
The use of GM applied to archaeological material culture has
seen a sharp increase in the past decade, yielding interesting results
regarding morphological variation among samples and generating
exciting knowledge about how such a variation can be explained in
terms of function, development or evolution. Given this scenario, it
would be expected that more researchers adopt GM to further
explore the morphology of all sorts of material culture. However,
this response has not been observed. Lycett (2009c, see also Lycett
and Chauhan, 2010) discusses why there were so few archaeolo-
gists working with GM (as well as other quantitative methods)
applied to lithic artefacts. The author offers three possible expla-
nations for that: lack of quantitative and statistical training, the
requirement of expensive digital equipment, and the difficulties of
using such instruments in archaeological field conditions. Other
less practical explanations include researchers feeling suspicious
about the “reality” of quantitative methods, as well as the idea that
the only feasible way of properly analysing an artefact has to
include information on technology. Specifically regarding GM
applied to Archaeology, there is the possibility that some archae-
ologists do not feel completely comfortable borrowing a method
from Biology, given the (mostly) not explicit assumptions that such
borrowing implies. That includes the potentially strong relation
between the adopted method (GM) and a given theory (Evolu-
tionary Archaeology). Although such a relation is not mandatory, it
is quite obvious that the common agenda shared by Evolutionary
Archaeology and Evolutionary Biology would facilitate the
borrowing of GM from the latter to the former discipline.
Another interesting observation is that the majority of GM
analysis in Archaeology has been focused on stone tools, mainly
projectile points (see Lycett and Chauhan, 2010 for a discussion
about the lack of easily defined points of homology on most stone
artefact forms). There can be some explanations for this bias,
including particularities in the morphology of such artefacts (for
 M. Okumura, A.G.M. Araujo / Journal of Archaeological Science xxx (2018) 1e10
example, presenting quite obvious points showing correspondence
across specimens, like blade tip, ears, and stem base), as well as
their quite flat morphology (Buchanan and Collard, 2010a), mean-
ing that a large component of the variation occurs in a single plane
(Velhagen and Roth, 1997) which allows the performance of two-
dimensional analysis (which are cheaper to do in comparison
with three-dimensional analysis). Another potential explanation
can be related to the previous knowledge regarding different point
morphologies associated with different chronological settings and
regional groups, which might have stimulated the use of GM to
further address such relations. It is quite possible that applying GM
to other kinds of material culture (like pottery, to cite one possi-
bility, see Martínez-Carrillo et al., 2010; Selden et al., 2014; Wilczek
et al., 2014) would result in interesting information and would
generate important knowledge about such artefacts.
Yet another important issue to be pursued in the GM studies of
material culture would be the recognition of important particu-
larities related to production techniques, as well as functional or
stylistic requirements. These idiosyncrasies include the impact of
reductive vs. additive technology (Schillinger et al., 2014a, 2014b,
2015), as well as raw material characteristics (Cardillo, 2010; Eren
et al., 2011, 2014), different levels of manufacturer skills (Eerkens
and Bettinger, 2001), among others. Such particularities might be
responsible for a greater range of variation within the same type of
artefacts in comparison with most living organisms and therefore,
explanations related to these trends of variation should be cali-
brated accordingly (Cardillo, 2010).
Previous discussions held by biologists regarding several
important GM concepts should not be ignored by archaeologists
who want to apply such method to material culture studies.
However, it has been over a decade since the first borrowings of GM
from Biology to Archaeology, so the time is ripe for archaeologists
to both increase their awareness about the proper adjustments that
should be made when adopting such a method and to prevent the
misuse or misinformed use of GM in archaeological studies.
Acknowledgements
The authors would like to thank Thilo Rehren, Christian Hog-
gard, and Sarah Stark for the assistance with the publication of this
article. We also would like to thank the two anonymous reviewers
for their insightful comments on the manuscript. Rolando
Gonz

and Judith Charlin kindly allowed us to use an
alez-Jose
adapted version of a figure from their work. This research was
financially supported by British Academy/Newton Mobility Grant
Scheme 2014 (MO, NG140077), the Brazilian National Council for
Scientific and Technological Development (MO, CNPq 303566/
2014-0, 443169/2014-9, 443242/2015-1), and the Sabbatical Pro-
gram at the Institute for Advanced Studies, University of Sa~o Paulo
(AGMA).
References
Adams, D.C., Rohlf, F.J., Slice, D.E., 2004. Geometric morphometrics: ten years of
progress following the ‘revolution’. Ital. J. Zool. 71, 5e16.
Ahler, S.A., Geib, P.R., 2000. Why flute? Folsom point design and adaptation.
J. Archaeol. Sci. 27, 799e820.
Andrews, B.N., Knell, E.J., Eren, M.I., 2015. The three lives of a uniface. J. Archaeol.
Sci. 54, 228e236.
Aneja, D., Vora, S.R., Camci, E.D., Shapiro, L.G., Cox, T.C., 2015. Automated detection
of 3D landmarks for the elimination of non-biological variation in geometric
morphometric analyses. In: 28th IEEE International Symposium on Computer-
based Medical Systems, pp. 78e83. Sa ~o Carlos.
Archer, W., Braun, D.R., 2010. Variability in bifacial technology at Elandsfontein,
Western cape, South Africa: a geometric morphometric approach. J. Archaeol.
Sci. 37, 201e209.
Barrientos, G., 2010. On the problem of identifying homologies in lithic artifacts. In:
Muscio, H., Cardillo, M. (Eds.), Argentine Archaeology and the Legacy of Charles
Please cite this article in press as: Okumura, M., Araujo, A.G.M., Archaeology, biology, and borrowing: A critical examination of Geometric
Morphometrics in Archaeology, Journal of Archaeological Science (2018), https://doi.org/10.1016/j.jas.2017.09.015
7
Darwin. British Archaeological Reports, Archaeopress, Oxford.
Beck, C., Jones, G.T., 2007. Early Paleoarchaic point morphology and chronology. In:
Graff, K., Schmidt, D.N. (Eds.), Paleoindian or Paleoarchaic? Great Basin Human
Ecology at the Pleistoceneeholocene Transition. University of Utah Press, Salt
Lake City, pp. 23e41.
Bettinger, R.L., Baumhoff, M.A., 1982. The Numic spread: great Basin cultures in
competition. Am. Antiq. 47, 485e503.
Binford, L.R., 1962. Archaeology as anthropology. Am. Antiq. 28, 217e225.
Binford, L.R., 1964. A consideration of archaeological research design. Am. Antiq. 29,
425e441.
Binford, L.R., 1965. Archaeological systematics and the study of culture process. Am.
Antiq. 31, 203e210.
Binford, L.R., 1987. Data, relativism and archaeological science. Man 22, 391e404.
Bleed, P., 1986. The optimal design of hunting weapons: maintainability or reli-
ability. Am. Antiq. 51, 737e747.
Bookstein, F.L., 1990. Introduction and overview. In: Rohlf, F.J., Bookstein, F.L. (Eds.),
Proceedings of the Michigan Morphometrics Workshop. University of Michigan
Museum of Zoology, Ann Arbor, pp. 61e74. Special Publication 2.
Bookstein, F.L., 1991. Morphometric Tools for Landmark Data: Geometry and
Biology. Cambridge University Press, Cambridge.
Bookstein, F.L., 1997. Landmark methods for forms without landmarks: morpho-
metrics of group differences in outline shape. Med. Image Anal. 1, 225e243.
Bookstein, F.L., 1998. A hundred years of morphometrics. Acta Zool. Acad. Sci. H. 44,
7e59.
Bookstein, F., Strauss, R., Humphries, J., Chernoff, B., Elder, R., Smith, G., 1982.
A comment upon the uses of Fourier methods in systematic. Syst. Zool. 31,
85e92.
Boyd, R., Richerson, P., 1985. Culture and the Evolutionary Process. University of
Chicago Press, Chicago.
Broughton, J.M., 2002. Prey spatial structure and behavior affect archaeological tests
of optimal foraging models: examples from the Emeryville Shellmound verte-
brate fauna. World Archaeol. 34, 60e83.
Buchanan, B., 2006. An analysis of Folsom projectile point resharpening using
quantitative comparisons of form and allometry. J. Archaeol. Sci. 33, 185e199.
Buchanan, B., Collard, M., 2007. Investigating the peopling of North America
through cladistic analyses of Early Paleoindian projectile points. J. Anthropol.
Archaeol. 26, 366e393.
Buchanan, B., Collard, M., 2008a. Phenetics, cladistics, and the search for the
Alaskan ancestors of the Paleoindians: a reassessment of relationships among
the Clovis, Nenana, and Denali archaeological complexes. J. Archaeol. Sci. 35,
1683e1694.
Buchanan, B., Collard, M., 2008b. Testing models of early Paleoindian colonization
and adaptation using cladistics. In: O'Brien, M.J. (Ed.), Cultural Transmission and
Archaeology: Issues and Case Studies. Society for American Archaeology Press,
Washington, DC, pp. 59e76.
Buchanan, B., Collard, M., 2010a. A geometric morphometrics-based assessment of
blade shape differences among Paleoindian projectile point types from western
North America. J. Archaeol. Sci. 37, 350e359.
Buchanan, B., Collard, M., 2010b. An assessment of the impact of resharpening on
Paleoindian projectile point blade shape using geometric morphometric tech-
niques. In: Lycett, S.J., Chauhan, P.R. (Eds.), New Perspectives on Old Stones:
Analytical Approaches to Paleolithic Technologies. Springer, New York,
pp. 255e273.
Buchanan, B., Johnson, E., Strauss, R.E., Lewis, P.J., 2007. A morphometric approach
to assessing late Paleoindian projectile point variability on the Southern High
Plains. Plains Anthropol. 52, 279e299.
Buchanan, B., O'Brien, M.J., Collard, M., 2014. Continent-wide or region-specific? A
geometric morphometrics-based assessment of variation in Clovis point shape.
Archaeol. Anthropol. Sci. 6, 145e162.
Buchanan, B., Eren, M.I., Boulanger, M.T., O'Brien, M.J., 2015. Size, shape, scars, and
spatial patterning: a quantitative assessment of late Pleistocene (Clovis) point
resharpening. J. Archaeol. Sci. Rep. 3, 11e21.
Byers, D.A., Ugan, A., 2005. Should we expect large game specialization in the late
Pleistocene? An optimal foraging perspective on early Paleoindian prey choice.
J. Archaeol. Sci. 32, 1624e1640.
Cardillo, M., 2006. Explorando la variacio
n en las morfologías líticas a partir de las

cnicas de an

te alisis de contornos. El caso de las puntas de proyectil del hol-
oceno medio-tardío de la Puna de Salta (San Antonio de los Cobres, Argentina).
Werken 7, 77e88.
Cardillo, M., 2009. Temporal trends in the morphometric variation of the lithic
projectile points during the Middle Holocene of Southern Andes (Puna region) -
a coevolutionary approach. In: Muscio, H.J., Lo
pez, G.E.J. (Eds.), Theoretical and
Methodological Issues in Evolutionary Archaeology: toward an Unified
Darwinian Paradigm. Archaeopress, Oxford, pp. 13e20.
Cardillo, M., 2010. Some applications of geometric morphometrics to archaeology.
In: Elewa, A.M.T. (Ed.), Morphometrics to Nonmorphometricians. Springer,
pp. 325e341.
Cardillo, M., Borrazzo, K., Charlin, J., 2016. Environment, space, and morphological
variation of projectile points in Patagonia (Southern South America). Quat. Int.
422, 44e56.
Castin~ eira, C., Cardillo, M., Chalin, J., Baeza, J., 2011. Ana
lisis de morfometría geo-
me
trica en puntas Cola de Pescado del Uruguay. Lat. Am. Antiq. 22, 335e358.
Castin~ eira, C., Charlin, J., Cardillo, M., Baeza, J., 2012. Exploring morphometric var-
iations in fishtail projectile points from Uruguay, pampa, and Patagonia. In:
Miotti, L., Salemme, M., Flegenheimer, N., Goebel, T. (Eds.), Southbond: Late
8 M. Okumura, A.G.M. Araujo / Journal of Archaeological Science xxx (2018) 1e10
Pleistocene Peopling of Latin America. Center for the Study of the First Amer-
icans. Department of Anthropology, Texas A&M University, College Station,
pp. 9e13.
Cavalli-Sforza, L.L., Feldman, M.W., 1981. Cultural transmission and evolution: a
quantitative approach. Princeton University Press, Princeton.
Charlin, J., Gonz
alez-Jose
, R., 2012. Size and shape variation in late holocene pro-
jectile points of Southern Patagonia: a geometric morphometric study. Am.
Antiq. 77, 221e242.
Charlin, J., Cardillo, M., Borrazzo, K., 2014. Spatial patterns in Late Holocene lithic
projectile point technology of Tierra del Fuego (southern South America):
assessing size and shape changes. World Archaeol. 46, 78e100.
Chauhan, P.R., 2003. An overview of the Siwalik Acheulian and reconsidering its
chronological relationship with the Soanian e a theoretical perspective.
Assemblage 7, 1e28.
Cheverud, J.M., 1982. Relationships among ontogenetic, static, and evolutionary
allometry. Am. J. Phys. Anthropol. 59, 139e149.
Cheverud, J.M., 1984. Quantitative genetics and developmental constraints on
evolution by selection. J. Theor. Biol. 110, 155e171.
Cock, A.G., 1966. Genetical aspects of metrical growth and form in animals. Q. Rev.
Biol. 41, 131e190.
Collard, M., 2010. Integrating anthropological genetics with cultural anthropology
and archaeology: new opportunities. J. Anthropol. Sci. 88, 239e242.
Collard, M., Shennan, S.J., 2000. Ethnogenesis versus phylogenesis in prehistoric
culture change: a case-study using European Neolithic pottery and biological
phylogenetic techniques. In: Renfrew, C., Boyle, K. (Eds.), Archaeogenetics: DNA
and the Population Prehistory of Europe. McDonald Institute for Archaeological
Research, Cambridge, pp. 89e97.
Collard, M., Shennan, S.J., Tehrani, J.J., 2006. Branching, blending, and the evolution
of cultural similarities and differences among human populations. Evol. Hum.
Behav. 27, 169e184.
Cosmides, L., Tooby, J., 1992. Cognitive adaptations for social exchange. In:
Barkow, J.H., Cosmides, L., Tooby, J. (Eds.), The Adapted Mind. Oxford University
Press, New York and Oxford, pp. 163e228.
Costa, A.G., 2010. A geometric morphometric assessment of plan shape in bone and
stone Acheulean bifaces from the Middle Pleistocene site of Castel di Guido,
Latium, Italy. In: Lycett, S.J., Chauhan, P.R. (Eds.), New Perspectives on Old
Stones: Analytical Approaches to Paleolithic Technologies. Springer, New York,
pp. 23e41.
Coward, F., Shennan, S., Colledge, S., Conolly, J., Collard, M., 2008. The spread of
Neolithic plant economies from the Near East to northwest Europe: a phylo-
genetic analysis. J. Archaeol. Sci. 35, 42e56.
Darroch, J.N., Mosimann, J.E., 1985. Canonical and principal components of shape.
Biometrika 72, 241e252.
Darwent, J., O'Brien, M.J., 2006. Using cladistics to construct lineages of projectile
points from northeastern Missouri. In: CP, L., O'Brien, M.J., Collard, M.,
Shennan, S.J. (Eds.), Mapping Our Ancestors: Phylogenetic Approaches in An-
thropology and Prehistory. Aldine, New York, pp. 185e208.
Darwin, C., 1859. (1991). On the Origin of Species by Means of Natural Selection.
Murray, London.
Davis, L.G., Bean, D.W., Nyers, A.J., Brauner, D.R., 2015. GLiMR: a GIS-based method
for the geometric morphometric analysis of artifacts. Lithic Technol. 40,
199e217.
de Azevedo, S., Nocera, A., Paschetta, C., Castillo, L., Gonzalez, M., Gonza
lez-Jose

, R.,
2011. Evaluating microevolutionary models for the early settlement of the New
World: the importance of recurrent gene flow with Asia. Am. J. Phys. Anthropol.
146, 116e129.
de Azevedo, S., Charlin, J., Gonza
lez-Jose

, R., 2014. Identifying design and reduction
effects on lithic projectile point shapes. J. Archaeol. Sci. 41, 297e307.
de Azevedo, S., Quinto-Sa
nchez, M., Paschetta, C., Gonz

, R., 2017. The first
alez-Jose
human settlement of the New World: a closer look at craniofacial variation and
evolution of early and late Holocene Native American groups. Quat. Int. 431
(Part B), 152e167.
Dibble, H.L., 1984. Interpreting typological variation of Middle Paleolithic scrapers:
function, style, or sequence of reduction. J. Field Archaeol. 11, 431e436.
Dunnell, R.C., 1980. Evolutionary theory and archaeology. Adv. Archaeol. Method
Theory 3, 35e99.
Dunnell, R.C., 1982. Science, Social Science, and common sense: the agonizing
dilemma of modern Archaeology. J. Anthropol. Res. 38, 1e25.
Dunnell, R.C., 1992. Is a Scientific Archaeology Possible? Metaarchaeology. Springer,
pp. 75e97.
Eerkens, J.W., Bettinger, R.L., 2001. Techniques for assessing standardization in
artifact assemblages: can we scale material variability? Am. Antiq. 66, 493e504.
Eerkens, J.W., Lipo, C.P., 2007. Cultural transmission theory and the archaeological
record: providing context to understanding variation and temporal changes in
material culture. J. Archaeol. Res. 15, 239e274.
Eerkens, J.W., Bettinger, R.L., McElreath, R., 2006. Cultural transmission, phyloge-
netics, and the archaeological record. In: Lipo, C.P., O'Brien, M.J., Collard, M.,
Shennan, S.J. (Eds.), Mapping Our Ancestors: Phylogenetic Approaches in An-
thropology and Prehistory. Aldine, New York, pp. 169e183.
Ehrlich, R., Pharr, R.B., Healy-Williams, N., 1983. Comments on the validity of Fourier
descriptors in systematics: a reply to Bookstein et al. Syst. Zool. 32, 202e206.
Eren, M.I., Lycett, S.J., Roos, C.I., Sampson, C.G., 2011. Toolstone constraints on
knapping skill: levallois reduction with two different raw materials. J. Archaeol.
Sci. 38, 2731e2739.
Eren, M.I., Roos, C.I., Story, B.A., von Cramon-Taubadel, N., Lycett, S.J., 2014. The role
Please cite this article in press as: Okumura, M., Araujo, A.G.M., Archaeology, biology, and borrowing: A critical examination of Geometric
Morphometrics in Archaeology, Journal of Archaeological Science (2018), https://doi.org/10.1016/j.jas.2017.09.015
of raw material differences in stone tool shape variation: an experimental
assessment. J. Archaeol. Sci. 49, 472e487.
Evans, L.T., 1984. Darwin's use of the analogy between artificial and natural selec-
tion. J. Hist. Biol. 17, 113e140.
Flenniken, J.J., Raymond, A.W., 1986. Morphological projectile point typology:
replication experimentation and technological analysis. Am. Antiq. 51, 603e614.
Fox, A.N., 2015. A study of Late Woodland projectile point typology in New York
using elliptical Fourier outline analysis. J. Archaeol. Sci. Rep. 4, 501e509.
Franco, N.V., Castro, A., Cardillo, M., Charlin, J., 2009. La importancia de las variables
morfolo
gicas, me
tricas y de microdesgaste para evaluar las diferencias en dis-
en~ os de puntas de proyectil bifaciales pedunculadas: un ejemplo del Sur de
Patagonia continental. Magallania (Chile), vol. 37, pp. 99e112.
Franklin, D., Oxnard, C.E., O'Higgins, P., Dadour, I., 2007. Sexual dimorphism in the
subadult mandible: quantification using geometric morphometrics. J. Forensic
Sci. 52, 6e10.
Franklin, D., Cardini, A., O'Higgins, P., Oxnard, C.E., Dadour, I., 2008. Mandibular
morphology as an indicator of human subadult age: geometric morphometric
approaches. Forensic Sci. Med. Pathol. 4, 91e99.
Friess, M., Baylac, M., 2003. Exploring artificial cranial deformation using elliptic
Fourier analysis of Procrustes aligned outlines. Am. J. Phys. Anthropol. 122,
11e22.
Futuyma, D.J., 2010. Evolutionary constraint and ecological consequences. Evolution
64, 1865e1884.
Galland, M., 2015. Le premier peuplement des Ame
riques: application de la mor-
phome
trie ge

ome
trique 3D a  la variation cr^
anienne actuelle et fossile. B Mem.
Soc. Anthro Par. 27, 189e201.
Galland, M., Friess, M., 2016. A three-dimensional geometric morphometrics view of
the cranial shape variation and population history in the New World. Am. J.
Hum. Biol. 28, 646e661.
Gero, J., Mazzullo, J., 1984. Analysis of artifact shape using Fourier series in closed
form. J. Field Archaeol. 11, 315e322.
Gonzalez, P.N., Bernal, V., Perez, S.I., 2009. Geometric morphometric approach to
sex estimation of human pelvis. Forensic Sci. Int. 189, 68e74.
Gonzalez, P., Bernal, V., Perez, S., 2011. Analysis of sexual dimorphism of craniofacial
traits using geometric morphometric techniques. Int. J. Osteoarchaeol 21,
82e91.
Gonz
alez-Jose
, R., Charlin, J., 2012. Relative importance of modularity and other
morphological attributes on different types of lithic point weapons: assessing
functional variations. PloS one 7, e48009.
Gonz
alez-Jose
, R., Gonz
alez-Martín, A., Hern
andez, M., Pucciarelli, H.M., Sardi, M.,
Rosales, A., Van der Molen, S., 2003. Craniometric evidence for palaeoamerican
survival in baja California. Nature 425, 62e65.
Gonz
alez-Jose
, R., Bortolini, M.C., Santos, F.R., Bonatto, S.L., 2008. The peopling of
America: craniofacial shape variation on a continental scale and its interpre-
tation from an interdisciplinary view. Am. J. Phys. Anthropol. 137, 175e187.
Gould, S.J., 1966. Allometry and size in ontogeny and phylogeny. Biol. Rev. 41,
587e638.
Gunz, P., Mitteroecker, P., Bookstein, F.L., 2005. Semilandmarks in three dimensions.
In: Slice, D.E. (Ed.), Modern Morphometrics in Physical Anthropology. Springer
US, New York, pp. 73e98.
Hall, B.K., 1994. Homology: the Hierarchial Basis of Comparative Biology. Academic
Press, San Diego. https://books.google.com.br/books?id¼LvzKBAAAQBAJ
&printsec¼frontcover&dq¼hallþhomologyþtheþhierarchicalþbasisþ1994&hl
¼en&sa¼X&ved¼0ahUKEwj0pOzYwMTaAhUFE5AKHScVAqUQ6AEIKTAA#v¼
onepage&q¼hall%20homology%20the%20hierarchical%20basis%201994&f¼
false.
Halloway Jr., R.L., 1969. Culture: a human domain. Curr. Anthropol. 10, 395e407.
Harmon, M.J., VanPool, T.L., Leonard, R.D., VanPool, C.S., Salter, L.A., 2006. Recon-
structing the flow of information across time and space: a phylogenetic analysis
of ceramic traditions from prehispanic western and northern Mexico and the
American Southwest. In: Lipo, C.P., O'Brien, M.J., Collard, M., Shennan, S.J. (Eds.),
Mapping Our Ancestors: Phylogenetic Approaches in Anthropology and Pre-
history. Aldine, New York, pp. 209e229.
Hayden, B., 1987. From chopper to celt: the evolution of resharpening techniques.
Lithic Technol. 16, 33e43.
Hoffman, C.M., 1985. Projectile point maintenance and typology: assessment with
factor analysis and canonical correlation. In: Carr, C. (Ed.), For Concordance in
Archaeological Analysis: Bridging Data Structure, Quantitative Technique, and
Theory. Westport Publishers, Kansas City, Mo. and Fayetteville, pp. 566e612.
Hofman, J.L., Graham, R.W., 1998. The paleo-indian cultures of the great plains. In:
Wood, W.R. (Ed.), Archaeology on the Great Plains. University Press of Kansas,
Lawrence, pp. 87e139.
Hunt, T.L., Lipo, C.P., Sterling, S.L., 2001. Posing Questions for a Scientific Archae-
ology. Bergin & Garvey, Westport and London.
Iovita, R., 2009. Ontogenetic scaling and lithic systematics: method and application.
J. Archaeol. Sci. 36, 1447e1457.
Iovita, R., 2010. Comparing stone tool resharpening trajectoires with the aid of
elliptical fourier analysis. In: Lycett, S.J., Chauhan, P.R. (Eds.), New Perspectives
on Old Stones: Analytical Approaches to Palaeolithic Technologies. Springer,
New York, pp. 235e253.
Iovita, R., 2011. Shape variation in Aterian tanged tools and the origins of projectile
technology: a morphometric perspective on stone tool function. PloS one 6,
e29029.
Iovita, R., McPherron, S.P., 2011. The handaxe reloaded: a morphometric reassess-
ment of Acheulian and Middle Paleolithic handaxes. J. Hum. Evol. 61, 61e74.
 M. Okumura, A.G.M. Araujo / Journal of Archaeological Science xxx (2018) 1e10
Jardine, N., 1969. The observational and theoretical components of homology: a
study based on the morphology of the dermal skull-roofs of rhipidistian fishes.
Biol. J. Linn. Soc. 1, 327e361.
Jonker, C.M., Snoep, J.L., Treur, J., Westerhoff, H.V., Wijngaards, W.C., 2002. Putting
intentions into cell biochemistry: an artificial intelligence perspective. J. Theor.
Biol. 214, 105e134.
Jordan, P., Shennan, S.J., 2003. Cultural transmission, language, and basketry tra-
ditions amongst the Californian Indians. J. Anthropol. Archaeol. 22, 42e74.
Keene, A.S., 1983. Biology, behavior, and borrowing: a critical examination of
Optimal Foraging Theory in archaeology. In: Moore, J.A., Keene, A.S. (Eds.),
Archaeological Hammers and Theories. Academic Press, New York.
Klingenberg, C.P., 1996. Multivariate allometry. In: Marcus, L.F., Corti, M., Loy, A.,
Naylor, G.J.P., Slice, D.E. (Eds.), Advances in Morphometrics. Springer, New York,
pp. 23e49.
Klingenberg, C.P., 1998. Heterochrony and allometry: the analysis of evolutionary
change in ontogeny. Biol. Rev. Camb Philos. 73, 79e123.
Klingenberg, C.P., 2008. Morphological integration and developmental modularity.
Annu. Rev. Ecol. Evol. S 39, 115e132.
Klingenberg, C.P., 2009. Morphometric integration and modularity in configurations
of landmarks: tools for evaluating a priori hypotheses. Evol. Dev. 11, 405e421.
Klingenberg, C.P., 2013. Cranial integration and modularity: insights into evolution
and development from morphometric data. Hystrix 24, 43e58.
Klingenberg, C.P., Zimmermann, M., 1992. Static, ontogenetic, and evolutionary
allometry: a multivariate comparison in nine species of water striders. Am. Nat.
140, 601e620.
Klingenberg, C.P., Badyaev, A.V., Sowry, S.M., Beckwith, N.J., 2001. Inferring devel-
opmental modularity from morphological integration: analysis of individual
variation and asymmetry in bumblebee wings. Am. Nat. 157, 11e23.
Kooyman, B.P., 2000. Understanding Stone Tools and Archaeological Sites. Univer-
sity of Calgary Press, Calgary.
Krohs, U., 2009. The cost of modularity. In: Krohs, U., Kroes, P. (Eds.), Functions in
Biological and Artificial Worlds: Comparative Philosophical Perspectives. MIT
Press, Cambridge, pp. 259e276.
Lawing, A.M., Polly, P.D., 2010. Geometric morphometrics: recent applications to the
study of evolution and development. J. Zool. 280, 1e7.
Lenardi, M.J., Merwin, D.E., 2010. Towards Automating Artifact Analysis: a Study
Showing Potential Applications of Computer Vision and Morphometrics to
Artifact Typology. Morphometrics for Nonmorphometricians. Springer, New
York, pp. 289e305.
Lycett, S.J., 2007. Why is there a lack of Mode 3 Levallois technologies in East Asia? A
phylogenetic test of the MoviuseSchick hypothesis. J. Anthropol. Archaeol. 26,
541e575.
Lycett, S.J., 2009a. Understanding ancient hominin dispersals using artefactual data:
a phylogeographic analysis of Acheulean handaxes. Plos One 4, e7404.
Lycett, S.J., 2009b. Are Victoria West cores ‘‘proto-Levallois’’? A phylogenetic
assessment. J. Hum. Evol. 56, 175e191.
Lycett, S.J., 2009c. Quantifying transitions: morphometric approaches to Palae-
olithic variability and technological change. In: Camps, M., Chauhan, P. (Eds.),
Sourcebook of Paleolithic Transitions. Springer, New York, pp. 79e92.
Lycett, S.J., 2010. Cultural transmission, genetic models and Palaeolithic variability:
integrative analytical approaches. In: Lycett, S., Chauhan, P. (Eds.), New Per-
spectives on Old Stones. Springer, New York, pp. 207e234.
Lycett, S.J., 2015. Cultural evolutionary approaches to artifact variation over time
and space: basis, progress, and prospects. J. Archaeol. Sci. 56, 21e31.
Lycett, S.J., Chauhan, P.R., 2010. Analytical approaches to Palaeolithic technologies:
an introduction. In: Lycett, S., Chauhan, P. (Eds.), New Perspectives on Old
Stones. Springer, New York, pp. 1e22.
Lycett, S.J., Von Cramon-Taubadel, N., 2012. A 3D morphometric analysis of surface
geometry in Levallois cores: patterns of stability and variability across regions
and their implications. J. Archaeol. Sci. 40, 1508e1517.
Lycett, S.J., von Cramon-Taubadel, N., 2013. A 3D morphometric analysis of surface
geometry in Levallois cores: patterns of stability and variability across regions
and their implications. J. Archaeol. Sci. 40, 1508e1517.
Lycett, S.J., Von Cramon-Taubadel, N., Foley, R.A., 2006. A crossbeam co-ordinate
caliper for the morphometric analysis of lithic nuclei: a description, test and
empirical examples of application. J. Archaeol. Sci. 33, 847e861.
Lycett, S.J., von Cramon-Taubadel, N., Gowlett, J.A.J., 2010. A comparative 3D geo-
metric morphometric analysis of Victoria West cores: implications for the ori-
gins of Levallois technology. J. Archaeol. Sci. 37, 1110e1117.
Lyman, R.L., O'Brien, M.J., Dunnell, R.C., 1997. The Rise and Fall of Culture History.
Plenum Press, New York.
MacLeod, N., 1999. Generalizing and extending the eigenshape method of shape
space visualization and analysis. Paleobiology 25, 107e138.
MacLeod, N., 2002. Phylogenetic signals in morphometric data. In: MacLeod, N.,
Forey, P.L. (Eds.), Morphology, Shape and Phylogeny. Taylor & Francis, London.
MacLeod, N., 2008. Understanding morphology in systematic contexts: 3D spec-
imen ordination and 3D specimen recognition. In: Wheeler, Q. (Ed.), The New
Taxonomy. CRC Press, Taylor & Francis Group, London, pp. 143e210.
Manríquez, G., Moraga, M., Santoro, C.M., Aspillaga, E., Arriaza, B.T., Rothhammer, F.,
2011. Morphometric and mtDNA analyses of archaic skeletal remains from
southwestern South America. Chungara 43, 283e292.
Martínez-Abadías, N., Gonza
lez-Jose

, R., Gonza

lez-Martín, A., Van der Molen, S.,
Talavera, A., Herna
ndez, P., Herna
ndez, M., 2006. Phenotypic evolution of hu-
man craniofacial morphology after admixture: a geometric morphometrics
approach. Am. J. Phys. Anthropol. 129, 387e398.
Please cite this article in press as: Okumura, M., Araujo, A.G.M., Archaeology, biology, and borrowing: A critical examination of Geometric
Morphometrics in Archaeology, Journal of Archaeological Science (2018), https://doi.org/10.1016/j.jas.2017.09.015
9
Martínez-Carrillo, A.L., Lucena, M.J., Fuertes, J.M., Ruiz, A., 2010. Morphometric
analysis applied to the archaeological pottery of the valley of Guadalquivir. In:
Elewa, A.M.T. (Ed.), Morphometrics for Nonmorphometricians. Springer, Berlin
and Heidelberg, pp. 307e323.
Marwick, B., 2012. A cladistic evaluation of ancient Thai bronze Buddha images: six
tests for a phylogenetic signal in the Griswold collection. In: Tjoa-Bonatz, M.L.,
Reinecke, A., Bonatz, D. (Eds.), Connecting Empires and States (Selected Papers
from the 13th International Conference of the European Association of South-
east Asian Archaeologists). NUS Press, Singapore, pp. 159e176.
Mayr, E., 1959. Darwin and the evolutionary theory in biology. In: Meggers, B.J. (Ed.),
Evolution and Anthropology: a Centennial Appraisal. The Anthropological So-
ciety of Washington, pp. 1e10.
McPherron, S.P., Dibble, H.L., 1999. Stone tool analysis using digitized images: ex-
amples from the lower and middle paleolithic. Lithic Technol. 24, 38e52.
Mitteroecker, P., Bookstein, F., 2007. The conceptual and statistical relationship
between modularity and morphological integration. Syst. Biol. 56, 818e836.
Mitteroecker, P., Gunz, P., 2009. Advances in geometric morphometrics. Evol. Biol.
36, 235e247.
Musil, R.R., 1988. Functional efficiency and technological change: a hafting tradition
model for prehistoric North America. In: Willig, J.A., Aikens, C.M., Fagan, J.L.
(Eds.), Early Human Occupation in Far Western North America: the Clovis-
Archaic Interface. Anthropological Papers No. 21. Nevada State Museum,
Nevada, pp. 373e387.
Neiman, F.D., 1995. Stylistic variation in evolutionary perspective. Ame Antiq. 60,
7e36.
Neves, W.A., Hubbe, M., Okumura, M.M.M., Gonz
alez-Jose
, R., Figuti, L., Eggers, S.,
DeBlasis, P.A.D., 2005. A new early Holocene human skeleton from Brazil: im-
plications for the settlement of the New World. J. Hum. Evol. 48, 403e414.
O'Brien, M.J., Darwent, J., Lyman, R.L., 2001. Cladistics is useful for reconstructing
archaeological phylogenies: palaeoindian points from the Southeastern United
States. J. Archaeol. Sci. 28, 1115e1136.
O'Brien, M.J., Lyman, R.L., Saab, Y., Saab, E., Darwent, J., Glover, D.S., 2002. Two issues
in archaeological phylogenetics: taxon construction and outgroup selection.
J. Theor. Biol. 215, 133e150.
O'Brien, M.J., Lyman, R.L., Mesoudi, A., VanPool, T., 2010. Cultural traits as units of
analysis. Philos. T R. Soc. B 365, 3797e3806.
O'Higgins, P., 2000. The study of morphological variation in the hominid fossil re-
cord: biology, landmarks and geometry. J. Anat. 197, 103e120.
Oestigaard, T., 2004. The world as artefact: material culture studies and Archae-
ology. In: Fahlander, F., Oestigaard, T. (Eds.), Material Culture and Other Things.
Post-disciplinary Studies in the 21st Century. Gotarc Series C 61. University of
Gothenburg, Gothenburg, pp. 21e55.
Okumura, M., Araujo, A.G.M., 2013. Pontas bifaciais no Brasil Meridional:
caracterizaça ~o estatística das formas e suas implicaço ~es culturais. Rev. do Mus.
Arqueol. Etnologia 23, 111e127.
Okumura, M., Araujo, A.G.M., 2014. Long-term cultural stability in hunter-
egatherers: a case study using traditional and geometric morphometric anal-
ysis of lithic stemmed bifacial points from Southern Brazil. J. Archaeol. Sci. 45,
59e71.
Okumura, M., Araujo, A.G.M., 2016. The Southern Divide: testing morphological
differences among bifacial points from southern and southeastern Brazil using
geometric morphometrics. J. Lithic Stud. 3, 107e132.
Olson, E.C., Miller, R.L., 1999. Morphological Integration. University of Chicago Press,
Chicago.
Owen, R., 1848. On the Archetype and Homologies of the Vertebrate Skeleton. van
Voorst, London.
O'Brien, M.J., Lyman, R.L., 2003. Cladistics and Archaeology. University of Utah Press,
Salt Lake City.
O'Brien, M.J., Buchanan, B., Collard, M., Boulanger, M.T., 2012. Cultural cladistics and
the early prehistory of North America. In: Pontarotti, P. (Ed.), Evolutionary
Biology: Mechanisms and Trends. Springer, Berlin, pp. 23e42.
Perez, S.I., 2007. Artificial cranial deformation in South America: a geometric
morphometrics approximation. J. Archaeol. Sci. 34, 1649e1658.
Perez, S.I., Bernal, V., Gonzalez, P.N., Sardi, M., Politis, G.G., 2009. Discrepancy be-
tween cranial and DNA data of early Americans: implications for American
peopling. PLoS One 4, e5746.
Picin, A., Vaquero, M., Weniger, G.C., Carbonell, E., 2014. Flake morphologies and
patterns of core configuration at the Abric Romaní rock-shelter: a geometric
morphometric approach. Quatern Int. 350, 84e93.
Pinhasi, R., von Cramon-Taubadel, N., 2009. Craniometric data supports demic
diffusion model for the spread of agriculture into Europe. PloS one 4, e6747.
Pomidor, B.J., Makedonska, J., Slice, D.E., 2016. A landmark-free method for three-
dimensional shape analysis. PloS one 11 e0150368.
Prentiss, A.M., Skelton, R.R., Eldredge, N., Quinn, C., 2011. Get rad! the evolution of
the skateboard deck. Evol. Educ. Outreach 4, 379e389.
Raab, L.M., 1992. An optimal foraging analysis of prehistoric shellfish collecting on
San Clemente Island, California. J. Ethnobiol. 12, 63e80.
Raff, R.A., Wolpert, L., 1996. The Shape of Life-genes, Development and Evolution of
Animal Forms. Cambridge University Press, London and New York.
Read, D.W., Lestrel, P.E., 1986. Comment on uses of homologous-point measures in
systematics: a reply to Bookstein et al. Syst. Zool. 35, 241e253.
Rivero, D.G., 2016. The evolution of anthropomorphism in the neolithic engraved
plaques of Southwestern Iberian Peninsula: a systematic approach from phy-
logenetics. Archaeol. Anthropol. Sci. 9, 1689.
Rivero, D.G., O'Brien, M.J., 2014. Phylogenetic analysis shows that Neolithic slate
10 M. Okumura, A.G.M. Araujo / Journal of Archaeological Science xxx (2018) 1e10
plaques from the southwestern Iberian Peninsula are not genealogical
recording systems. PloS one 9, e88296.
Rohlf, F.J., 1986. Relationships among eigenshape analysis, Fourier analysis, and
analysis of coordinates. Math. Geol. 18, 845e854.
Rorabaugh, A.N., 2012. Prestige, transmission, and barbed bone and antler points in
the Gulf of Georgia, Northwest Coast. J. Contemp. Anthropol. 3, 17e37.
Sandstrom, G., 2013. Peace for evolution's puzzle: the arrival of human extension.
In: Grinin, L.E., Korotaiev, A.V. (Eds.), Evolution: Development within Big His-
tory, Evolutionary, and World-system Paradigms. Russian Academy of Sciences,
Volgograd Center for Social Research, Volgograd, pp. 267e288.
Scartascini, F.L., Cardillo, M., Palacios, T., Palacios, O., V
azquez, C., Cabanillas, E.,
2009. Explorando la variabilidad me
trica y morfolo
gica de las “pesas líticas”
recuperadas en el sector norte de la costa del golfo San Matías. Arqueometría
Latinoamericana: Segundo Congreso Argentino y Primero Latinoamericano.
Comisio
n Nacional de Energía Ato
mica, Buenos Aires, pp. 162e168.
Schiffer, M.B., 1972. Archaeological context and systemic context. Am. Antiq. 37,
156e165.
Schillinger, K., Mesoudi, A., Lycett, S., 2014a. Copying error and the cultural evolu-
tion of “additive” vs. “reductive” material traditions: an experimental assess-
ment. Am. Antiq. 79, 128e143.
Schillinger, K., Mesoudi, A., Lycett, S.J., 2014b. Considering the role of time budgets
on copy-error rates in material culture traditions: an experimental assessment.
PloS one 9 (5), e97157.
Schillinger, K., Mesoudi, A., Lycett, S.J., 2015. The impact of imitative versus
emulative learning mechanisms on artifactual variation: implications for the
evolution of material culture. Evol. Hum. Behav. 36, 446e455.
Schlosser, G., Wagner, G.P., 2004. The modularity concept in developmental and
evolutionary biology. In: Schlosser, G., Wagner, G.P. (Eds.), Modularity in
Development and Evolution. University of Chicago Press, Chicago and London,
pp. 1e11.
Selden Jr., R.Z., Perttula, T.K., O'Brien, M.J., 2014. Advances in documentation, digital
curation, virtual exhibition, and a test of 3D geometric morphometrics: a case
study of the Vanderpool vessels from the ancestral Caddo territory. Adv.
Archaeol. Pract. 2, 64e79.
Service, E.R., 1969. Models for the methodology of mouthtalk. Southwest J. Anthr.
25, 68e80.
Serwatka, K., 2015a. Shape variation of Middle Palaeolithic bifacial tools from
southern Poland: a geometric morphometric approach to Keilmessergruppen
handaxes and backed knives. Lithic J. Lithic Stud. Soc. 35, 18e32.
Serwatka, K., 2015b. Bifaces in plain sight: testing elliptical Fourier analysis in
identifying reduction effects on Late Middle Palaeolithic bifacial tools. Litikum
3, 13e25.
Shennan, S.J., 2000. Population, culture history, and the dynamics of culture change.
Curr. Anthropol. 41, 811e835.
Shennan, S.J., 2001. Demography and cultural innovation: a model and its impli-
cations for the emergence of modern human culture. Camb Archaeol. J. 11, 5e16.
Sherif, M., Sherif, C.W., 2009. Interdisciplinary coordination as a validity check:
retrospect and prospects. In: Sherif, M., Sherif, C.W. (Eds.), Interdisciplinary
Relationships in the Social Sciences. Transaction Publishers, New Brunswick
and London, pp. 3e20.
Shott, M.J., 2005. The reduction thesis and its discontents: review of Australian
approaches. In: Clarkson, C., Lamb, L. (Eds.), Lithics ‘Down under’: Australian
Perspectives on Lithic Reduction, Use and Classification. Archaeopress, Oxford,
pp. 109e125.
Shott, M.J., 2010. Stone-tool demography: reduction distributions in North Amer-
ican Paleoindian tools. In: Lycett, S., Chauhan, P. (Eds.), New Perspectives on Old
Stones. Springer, New York, pp. 275e293.
Shott, M.J., Ballenger, J.A., 2007. Biface reduction and the measurement of Dalton
curation: a southeastern United States case study. Am. Antiq. 72, 153e175.
Please cite this article in press as: Okumura, M., Araujo, A.G.M., Archaeology, biology, and borrowing: A critical examination of Geometric
Morphometrics in Archaeology, Journal of Archaeological Science (2018), https://doi.org/10.1016/j.jas.2017.09.015
Shott, M.J., Trail, B.W., 2010. Exploring new approaches to lithic analysis: laser
scanning and geometric morphometrics. Lithic Technol. 35, 195e220. https://
www.tandfonline.com/doi/abs/10.1080/01977261.2010.11721090.
Shott, M.J., Hunzicker, D.A., Patten, B., 2007. Pattern and allometric measurement of
reduction in experimental Folsom bifaces. Lithic Technol. 32, 203e217.
Simon, H.A., 1962. The architecture of complexity. P Am. Philos. Soc. 106, 467e482.
Slice, D.E., 2005. Modern morphometrics. In: Slice, D.E. (Ed.), Modern Morpho-
metrics in Physical Anthropology. Kluwer Academic/Plenun Publishers, New
York, pp. 1e45.
Smith, E.A., Winterhalder, B., 1981. Hunter-gatherer Foraging Strategies: Ethno-
graphic and Archeological Analysis. University of Chicago Press, Chicago.
Tehrani, J.J., Collard, M., 2002. Investigating cultural evolution through biological
phylogenetic analyses of Turkmen textiles. J. Anthropol. Archaeol. 21, 443e463.
Tehrani, J.J., Collard, M., 2009. An integrated analysis of inter-individual and inter-
group cultural transmission in Iranian tribal populations. Evol. Hum. Behav.
30, 286e300.
Temkin, I., Eldredge, N., 2007. Phylogenetics and material culture evolution. Curr.
Anthropol. 48, 146e153.
Thompson, D.W., 1942. On Growth and Form. Cambridge University Press,
Cambridge.
Thulman, D.K., 2012. Discriminating Paleoindian point types from Florida using
landmark geometric morphometrics. J. Archaeol. Sci. 39, 1599e1607.
Titmus, G.L., Woods, J.C., 1991. Fluted points from the snake river plain. In:
Bonnichsen, R., Turnmire, K.L. (Eds.), Clovis: Origins and Adaptations. Center for
the Study of the First Americans, Corvallis, pp. 119e131.
Tolstoy, P., 2008. Barkcloth, Polynesia and cladistics: an update. J. Polyn. Soc. 117,
15e57.
Towner, R.H., Warburton, M., 1990. Projectile point rejuvenation: a technological
analysis. J. Field Archaeol. 17, 311e321.
Velhagen, W.A., Roth, V.L., 1997. Scaling of the mandible in squirrels. J. Morphol.
232, 107e132.
Viscosi, V., Cardini, A., 2011. Leaf morphology, taxonomy and geometric morpho-
metrics: a simplified protocol for beginners. PLoS One 6, e25630.
von Cramon-Taubadel, N., Pinhasi, R., 2011. Craniometric data support a mosaic
model of demic and cultural Neolithic diffusion to outlying regions of Europe.
Proc. Roy. Soc. B-Biol. Sci. rspb20102678.
Wagner, G.P., 1994. Homology and the mechanisms of development. In: Hall, B.K.
(Ed.), Homology: the Hierarchical Basis of Comparative Biology. Academic Press,
San Diego, pp. 273e299.
Wagner, G.P., Altenberg, L., 1996. Perspective: complex adaptations and the evolu-
tion of evolvability. Evolution 50, 967e976.
Wallerstein, I., 2003. Anthropology, Sociology, and other dubious disciplines. Curr.
Anthropol. 44, 453e465.
Wang, W., Lycett, S.J., Von Cramon-Taubadel, N., Jin, J.J.H., Bae, C.J., 2012. Compar-
ison of handaxes from Bose Basin (China) and the Western Acheulean indicates
convergence of form, not cognitive differences. PloS one 7, e35804.
Watson, P.J., LeBlanc, S.A., Redman, C.L., 1971. Explanation in Archaeology: an
Explicitly Scientific Approach. Columbia University Press, New York.
Watson, P.J., LeBlanc, S.A., Redman, C.L.R., 1984. Archeological Explanation: the
Scientific Method in Archeology. Columbia University Press, New York.
Wilczek, J., Monna, F., Barral, P., Burlet, L., Chateau, C., Navarro, N., 2014. Morpho-
metrics of Second Iron Age ceramicsestrengths, weaknesses, and comparison
with traditional typology. J. Archaeol. Sci. 50, 39e50.
Zelditch, M.L., Fink, W.L., Swiderski, D.L., 1995. Morphometrics, homology, and
phylogenetics: quantified characters as synapomorphies. Syst. Biol. 44,
179e189.
Zelditch, M.L., Swiderski, D.L., Sheets, D.H., Fink, W.L., 2004. Geometric Morpho-
metrics for Biologists: a Primer. Elsevier, Academic Press, New York.