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Joel D. Irish
Department of Anthropology,
University of Alaska Fairbanks,
Fairbanks, Alaska
99775-7720, U.S.A.
Received 13 January 2000
Revision received 29 March
2000 and accepted 19 May
2000
Keywords:Late Pleistocene,
Africa, Maghreb, Nubia,
dental anthropology,
morphological variation,
biological affinity estimation.
The Iberomaurusian enigma: North
African progenitor or dead end?
Data obtained during an ongoing dental investigation of African
populations address two long-standing, hotly debated questions.
First, was there genetic continuity between Late Pleistocene Ibero-
maurusians and later northwest Africans (e.g., Capsians, Berbers,
Guanche)? Second, were skeletally-robust Iberomaurusians and
northeast African Nubians variants of the same population? Ibero-
maurusians from Taforalt in Morocco and Afalou-Bou-Rhummel in
Algeria, Nubians from Jebel Sahaba in Sudan, post-Pleistocene
Capsians from Algeria and Tunisia, and a series of other samples were
statistically compared using 29 discrete dental traits to help estimate
diachronic local and regional affinities. Results revealed: (1) a rela-
tionship between the Iberomaurusians, particularly those from
Taforalt, and later Maghreb and other North African samples, and
(2) a divergence among contemporaneous Iberomaurusians and
Nubian samples. Thus, some measure of long-term population
continuity in the Maghreb and surrounding region is supported,
whereas greater North African population heterogenity during the
Late Pleistocene is implied.
2000 Academic Press
Journal of Human Evolution (2000) 39, 393–410
doi:10.1006/jhev.2000.0430
Available online at http://www.idealibrary.com on
Introduction
There are, among others, two important
questions pertaining to North African pre-
history that have been sources of bio-
archaeological contention for decades. First,
were Late Pleistocene Iberomaurusian
populations ancestral to subsequent north-
west Africans, or were they essentially a
dead-end, playing only a minor role in the
latter’s genetic make-up? Second, were the
Iberomaurusians closely related to contem-
poraneous Nubians of northeast Africa,
or are reported physical and cultural
similarities between populations merely
superficial?
The term Iberomaurusian refers to: (1) a
Late Paleolithic tool industry characterized
by microlithic backed, partially backed
obtuse-ended, and other bladelets (Camps,
1974;Close, 1977;Lubell et al., 1984), and
(2) the makers of this industry; both were
present in numerous northwest African
Maghreb sites (Afalou-Bou-Rhummel, La
Mouillah, Taza Cave I, Taforalt, etc.)
between 20,000years ago and the termi-
nal Pleistocene (Ferembach, 1962,1985;
Chamla, 1973,1975,1978;Camps, 1974;
Hachi, 1996,1997;Medig et al., 1996;
Lubell, 2000). Most sites are clustered along
the Maghreb littoral in caves and rock shel-
ters: several contain human burials (Lubell,
2000). In the past, Iberomarusian peoples
have been called Mechta-Afalou, Mechta
el-Arbi, and/or Mechtoid types (see
Ferembach, 1962,1985;Vallois, 1969;
Chamla, 1973,1975,1978;Camps, 1974;
Dutour, 1985). They are a skeletally-robust
population that resembles European
Cro-magnon to some extent (Briggs, 1955;
Chamla, 1978;Ferembach, 1962,1985;
Correspondence to: Dr Joel D. Irish, Department of
Anthropology, P.O. Box 757720, University of Alaska
Fairbanks, Fairbanks, AK 99775–7720, U.S.A. Tel.:
(907) 474-6755; E-mail: ffjdi@uaf.edu
0047–2484/00/100393+ 18$35.00/0 2000 Academic Press
Clark, 1989;Groves & Thorne, 1999),
although they are said to be more rugged
(Pond, 1928;Hiernaux, 1975) or to vary in
additional ways (Briggs, 1954;Ferembach,
1962;Vallois, 1969). The origin of Ibero-
maurusians is unresolved; they may have
come from Europe, West Asia, elsewhere in
Africa, or they may have evolved in situ in
North Africa (see Vallois, 1969;Camps,
1974;Hiernaux, 1975;Ferembach, 1962,
1985;Petit-Marie & Dutour, 1987;Henke,
1990;Bermudez de Castro, 1991).
Regarding the question of population
continuity, several workers (e.g., Balout,
1955;Chamla, 1973,1975,1978;Camps,
1974;Hiernaux, 1975) believe Iberomauru-
sians contributed little to the genetic
make-up of later northwest Africans, includ-
ing Berbers and Canary Island Guanches,
although a few maintain they provided input
to the latter’s gene pool (Vallois, 1969;
Ferembach, 1985). Whichever the case, all
of these researchers agree Iberomaurusian
peoples were replaced in the Maghreb by
skeletally-gracile Capsians in the terminal
Late Pleistocene; it is Capsians, then, who
were the immediate ancestors of Berbers
and others (e.g., Tauregs and, perhaps,
Toubou) (Camps, 1974;Chamla, 1973,
1975,1978;Chabeuf, 1975;Hiernaux,
1975). These manufacturers of backed
blades and endscrapers on blades or large
flakes, and microliths from light bladelets
(Camps, 1974;Lubell et al., 1984), have
historically been sub-divided into three,
essentially time-successive, cultural manifes-
tations: the Typical, Upper, and Neolithic of
Capsian Traditions (Vaufrey, 1933;Wulsin,
1941;Sheppard, 1987;Phillipson, 1994).
Evidence for intrapopulation skeletal diver-
sity has also been reported (see Chamla,
1973,1975,1978;Camps, 1974). Sites may
be open-air or occur within caves and rock
shelters; all contain many land snails shells,
and other diagnostic food and material
remains (Haverkort & Lubell, 1999;Lubell,
2000). A few researchers maintain that the
Capsians were indigenous (see below), but
others (Briggs, 1954,1955;Ferembach,
1962,1985:396; Camps, 1974;Chamla,
1978) believe these ‘‘proto-Mediterranean’’
peoples migrated into the area from the
east—perhaps from as far afield as West
Asia.
Conversely, craniometric and lithic analy-
ses by Lubell and co-workers (Lubell et al.,
1984;Sheppard & Lubell, 1990;Lubell,
2000) suggest there is continuity from
Iberomaurusian through Capsian times.
These researchers do not perceive a biocul-
tural hiatus, although there may have been
spatial divergence (Lubell, 1984,2000).
Some level of continuity has also been
implied in other studies (e.g., Cluzel, 1973;
Close, 1986;Hachi, 1996,1997;Ighilarhriz,
1996). Iberomaurusians may have even
survived into the early Holocene in the
Malinese Sahara (Petit-Maire & Dutour,
1987) and along the Atlantic coast
(Ferembach, 1985), and there are reports
(Balout & Briggs, 1949;Briggs, 1954,
1955;Ferembach, 1962;Chamla, 1973,
1975,1978;Camps, 1974;Camps-Fabrer,
1975) of robust Mechta-Afalou- or
Mechtoid-like skeletal remains in Capsian
and later Maghreb sites [e.g. Mechta
el-Arbi, Medjez II, Grotte des Hye`nes, Rio
Salado (see Camps, 1974 for a full listing)].
Therefore, they could indeed be ancestors to
later northwest Africans, including Cap-
sians. Moreover, Lubell et al. (1984) feel
that if there was eastern influence, it prob-
ably came from the Nile Valley (i.e., Late
Pleistocene Nubians) rather than West Asia.
Concerning the question of affinity to
contemporaneous peoples, previous studies
show that Late Pleistocene Nubians share a
number of similarities with Iberomauru-
sians; the former’s Qadan (ca. 15,000–
11,000 BP) microlithic industry was like
that in the Maghreb (Clark, 1970;
Ferembach, 1985;Phillipson, 1994), and
they had comparable burial practices—
as evident at the Jebel Sahaba cemetery
394 ..
(SMU 117) in Lower Nubia (Wendorf,
1968). In addition, skeletal similitude
between populations has been suggested
(Anderson, 1968;Clark, 1970;Greene &
Armelagos, 1972;Dutour, 1995;Lahr &
Arensburg, 1995). Iberomaurusian skel-
etons from Dar-es-Soltan and Taforalt in
Morocco, and Ali-Bacha and Afalou-
Bou-Rhummel in Algeria, are said to look
like 12,000 year-old Nubian remains from
Wadi Halfa and Jebel Sahaba, based on
craniometric and nonmetric research
(Anderson, 1968;Greene & Armelagos,
1972). These populations, in turn, re-
semble the Nazlet Khater (Thoma, 1984;
Dutour, 1995) and 25,000-year-old Wadi
Kubbaniya remains from Egypt (Angel &
Kelley, 1986;Wendorf & Schild, 1986).
Overall, crania in these populations are said
to share many features, including prominent
brow ridges, low rectangular orbits, project-
ing zygomatic arches, alveolar prognathism,
large facial foramina, low broad mandibular
rami, gonial eversion, and large complex
teeth (see Anderson, 1968;Greene, 1972;
Greene & Armelagos, 1972).
However, Bermudez de Castro’s (1991)
dental comparison between the small Wadi
Halfa Nubian sample and the Taforalt and
Afalou-Bou-Rhummel Iberomaurusians
was inconclusive. Further, Camps (1974)
reports that the Qadan tool industry differs
noticeably from that of the Iberomauru-
sians, and describes several physical differ-
ences between the two contemporaneous
populations. Along these lines, craniofacial
(Franciscus, 1995, personal communi-
cation, 1995), dental (Irish, 1999), and
post-cranial (Holliday, 1995) comparative
analyses of the Taforalt and Afalou-Bou-
Rhummel samples with Jebel Sahaba
Nubians have cast doubts on a close biologi-
cal affinity; all show that the former two
samples exbibit many features reminiscent
of later North Africans, whereas Late Pleis-
tocene Nubians are more like recent sub-
Saharan Africans. These findings are largely
supported by Groves & Thorne’s (1999)
recent cranial analyses.
With this review of purported population
origins and relationships as a backdrop, the
goal of the present study is to address
the two dichotomous questions posed at the
outset. For this purpose, I will use dental
analyses of 16 Late Pleistocene through
recent North African samples (n=818 indi-
viduals). This study comprises a morpho-
logical examination of two Iberomaurusian
samples, and a multivariate statistical com-
parison of these data with those in: (1)
indigenous northwest Africans, including
Capsians, Shawia and Kabyle Berbers, and
Guanches, (2) Late Pleistocene Jebel
Sahaba Nubians, (3) post-Pleistocene
Nubians and Egyptians, and (4) West
Asian-derived Carthaginians and Bedouin
Arabs from the Maghreb. The latter two sets
of nine samples are included to help facili-
tate a better understanding of North African
population history in a broader, more
region- and circum-Mediterranean-oriented
perspective.
Materials
The two Iberomaurusian samples (Figure
1), comprising 90 dentitions, were recorded
at the Institut de Pale´ontologie Humaine.
They were recovered from the cave site of
Taforalt (n=42) (abbreviated at TAF in
Figures 2 and 4) in Morocco, and the
Afalou-Bou-Rhummel (n=48) (AFA) rock
shelter in Algeria. Remains from these sites
were dated 16,750 BP and ca. 12,500–
10,500 BP, respectively (Chamla, 1978;
Vallois, 1969); however, Lubell et al. (1992)
recently suggested Taforalt is younger, and
Hachi’s (1996) re-dating of Afalou yielded
an older range of 13,120370 to 11,450
230 BP.
Capsian (CAP) remains are located at
the University of Minnesota, University
of Alberta, and Institut de Pale´ontologie
Humaine. Although background records are
395
incomplete, this small, heterogenous sample
comes from both Algerian and Tunisian
sites [including Mechta el-Arbi, Mechta-
Chateaudun, Aı¨n Dokkara, and Grotte des
Hye`nes (Camps, 1974;Chamla, 1975)],
which are apparently associated with the
Typical (n=2 inds), Upper (n=12), and
Neolithic of Capsian Tradition (n=8) indus-
tries. Inclusive Capsian dates generally
range from ca. 8500 to 5000 BP (Chamla,
1973;Camps, 1974;Sheppard, 1987),
though Lubell (1984,2000, personal com-
munication, 2000; Lubell et al., 1992) notes
that two sites, Aı¨n Misteheyia and Kef
Zoura D, date to ca. 9800 BP.
The Shawia Berber (SHA) sample con-
sists of 26 historic individuals who originally
lived just south of Constantine, Algeria
(Figure 1). The sample is curated at the
Muse´edel’Homme. Greenberg (1966)
characterizes Berbers as speaking one of
several dialects (e.g., Shawia) of a native
Berber language, which reflects an influence
from Phoenician, Latin, and Arabic sources
(Bynon, 1970). Such heterogeneity is con-
sistent with the fact that Berbers, especially
those from less mountainous regions in
Algeria and Morocco, like the Shawia,
exhibit signs of admixture with Arabs and
other nonindigenous groups (Carthaginian,
Greek, Roman, Spanish, Turkish, French)
(Wysner, 1945).
The Kabyle Berber (KAB) sample is
made up of 32 historic crania from the
Djurdjura Mountains of north Algeria
(Wysner, 1945). They are also curated at
the Muse´edel’Homme. Unlike many
Berbers, the Kabyle remained isolated
from the many outsiders who successively
conquered lands throughout North Africa
beginning roughly 2800 years ago; as such,
they experienced little genetic admixture
(Wysner, 1945).
The Canary Island Guanche (CAN)
sample consists of 163 crania, curated at the
Muse´edel’Homme, the American Museum
of Natural History, and the National
Museum of Natural History. Many
researchers believe they were closely related
to northwest African Berbers, and may have
been recipients of some gene flow from
Arabs and Carthaginians (Schwidetsky,
Figure 1. Origin locations of the 16 North African dental samples.
396 ..
1963;Mercer, 1980;Bermudez de Castro,
1989;Irish, 1993,1998b). The exact date of
the series is unknown, but radiocarbon
dating in grottos, caves, and tumuli, like
those from which the present remains
were removed, range between 2020 and
31070 BP, with a median age of 1600–
1100 years BP (Mercer, 1980;Bermudez de
Castro, 1989).
The late Paleolithic Nubian (NUB)
sample, ca. 14,500–12,500 BP, consists of
67 crania from Jebel Sahaba (and Tushka)
(see Figure 1) in Lower Nubia. Remains
were excavated from three late Paleolithic
cemeteries: SMU 67/5A, 80, and 117
(Wendorf, 1968); they are curated at
Southern Methodist University. As noted,
the remains are associated with a Qadan
microlithic industry level of technology,
estimated to have begun around 15,000 BP.
The other seven northeast African
samples are, as noted, included in the analy-
sis to help delineate Iberomaurusian affini-
ties at a regional level. Three samples com-
prise 12th Dynasty through Byzantine
Egyptians (4000–1400 BP) (Elliot Smith &
Wood-Jones, 1910;Baines & Malek, 1982)
from Lisht (LIS) (n=61), El Hesa (HES)
(n=72), and Kharga Oasis (KHA) (n=26).
They are at the American and National
Museums of Natural History. Egyptians
may be indigenous, or may have originated
in West Asia and migrated into Africa dur-
ing the Neolithic (Curto, 1972;Mourant,
1983). Whichever the case, by the Dynastic
period they represented a heterogeneous
people, from combining of many ethnic
elements (e.g., Mediterranean, Berber,
Nubian) (Curto, 1972;Davidson, 1974).
The other four samples are from Nubia.
One consists of 18th Dynasty Pharonic
Nubians (3575–3380 BP) (Trigger, 1976)
from Soleb (SOL) (n=32); the rest include
Meroitic (MER) (n=91), X-Group (XGR)
(n=39), and Christian (CHR) (n=18)
Nubians (2100–600 BP) from Semna
(Zabkar & Zabkar, 1982). The Pharonic
sample was recorded at the Muse´ede
l’Homme; the others are at Arizona State
University. Post-Pleistocene Nubian origins
are unclear; they may represent indigen-
ous peoples that possess a sub-Saharan
component (Greene, 1967,1972;Carlson &
Van Gerven, 1979), or heavily admixed
migrants (Irish & Turner, 1990;Turner &
Markowitz, 1990).
Lastly, the Carthaginian (CAR) and
Bedouin Arab (BED) samples are not indig-
enous to Africa; they are included to provide
some measure of comparison to West Asian
populations, which from a dental analysis
perspective is important though, to date,
rarely studied (see below). They are also of
interest because of reported admixture with
Berber and Guanche populations. The
Carthaginian sample consists of 28 crania
from Carthage, Tunisia (Figure 1). Twenty-
four were recovered from Punic levels
(Charles-Picard & Picard, 1969). The
remainder may date to Punic or Roman
times. All are curated at the Muse´ede
l’Homme in Paris. Carthage was founded
in 751 BC by Phoenicians, a West Asian
people from Lebanon (Charles-Picard &
Picard, 1969). Romans conquered Carthage
in 146 BC. The Bedouin Arab sample
(n=49) comprises a mix of historic crania:
36 from Morocco, 10 from Algeria, two
from Tunisia, and one from Libya. The
latter was recorded at the University of
Minnesota; the others are at the Muse´ede
l’Homme. Arabs first entered Africa along
the Suez isthmus in the seventh century.
More arrived in the 11th century, when
Bedouins immigrated from Syria (Julian,
1970;Hiernaux, 1975). They appear similar
to Berbers, with whom they are heavily
admixed (Julien, 1970;Hiernaux, 1975).
Methods
The present study is concerned with mor-
phological variation in the permanent den-
tition. Up to 36 nonmetric dental and
397
osseous oral traits were recorded in each
individual (see list in Table 1). However, the
maximum trait number for Iberomaurusians
and many Capsians was often less, due
to UI1 evulsion (detailed in Briggs, 1955;
Ferembach, 1962;Camps, 1974;Camps-
Fabrer, 1975;Caillard, 1978;Chamla,
1978;Medig et al., 1996) and extreme
attrition.
Except for midline diastema, each dental
feature is found in the Arizona State Univer-
sity (ASU) Dental Anthropology System
(Turner et al., 1991). System procedures are
based on well-established criteria for scoring
intratrait variation. Traits are recorded using
23 rank-scale reference plaques, that stand-
ardize scoring by providing representations
of minimum, maximum, and the most
common intermediate expressions (Turner
et al., 1991).
The reasons for selecting these traits are
described fully elsewhere (Irish, 1993,
1998a). In brief, they are easy to record,
resist wear, possess a high genetic compo-
nent in expression (Scott, 1973; Turner,
personal communication, 1986; Turner
et al., 1991;Scott & Turner, 1997), and as
Turner et al. (1991:13) state,
‘‘. . . the fossil record has shown that (whatever
their adaptive value) they evolve very slowly
and, altogether, . . . powerfully characterize
populations for affinity studies.’’
Along these lines, individual and collec-
tive expressions of the 36 traits are docu-
mented from elsewhere in Africa (Irish,
1993,1997,1998a,b,c;Johnson & Lovell,
1994), as well as Asia, Europe, India, and
the New World (Dahlberg, 1963;Hanihara,
1967; Turner, 1985b,1992a,b;Lukacs
et al., 1998). Such documentation facilitates
the future comparisons of suites of features
within and among world samples and
regions (see Scott & Turner, 1997;Irish,
1998c).
After selection of traits, a decision regard-
ing which antimere to score is required. One
common method entails counting the right
or left side in all individuals (Haeussler et al.,
1988). A second method is to score both
antimeres and then, allowing for possible
asymmetry, count the side that shows
highest expression (Turner & Scott, 1977).
Thus, if a grade 1 is scored for Bushman
canine on one antimere and a grade 0 is
present on the other, the grade 1 is used for
analysis. To maximize sample size if just one
side is present, that side is scored and
assumed to represent highest expression.
This standard ASU System protocol is
termed the individual counting procedure,
and assumes scoring for the individual’s
maximum genetic potential for that trait
(Turner, 1985a); it is used in the present
study.
Finally, due to a documented lack of trait
sexual dimorphism (Scott, 1973,1980;
Smith & Shegev, 1988;Bermudez de
Castro, 1989;Turner et al., 1991;Hanihara,
1992;Irish, 1993), it is standard System
procedure to pool the sexes (Irish, 1997);
this protocol is followed here. For a com-
plete description of ASU System pro-
cedures, traits, and rationale, see Turner
et al. (1991) and Scott & Turner (1997).
After recording, trait frequencies were
determined for each sample and C. A. B.
Smith’s Mean Measure of Divergence
(MMD) distance statistic, using the
Freeman and Tukey angular transformation
correction (Berry & Berry, 1967;Sjøvold,
1973;Green & Suchey, 1976) for small
sample sizes, and low (c0·05) or high
(d0·95) trait frequencies (Sjøvold, 1977)
was used. This multivariate technique pro-
vides a quantitative estimate of biological
divergence among samples based on the
degree of phenetic similarity for all traits. An
MMD calculated between sample pairs is a
dissimilarity measure; thus, lower values
indicate greater affinity, and vice versa. It is
assumed that phenetic similarity approxi-
mates or is an estimate of genetic variation
(Scott et al., 1983). To detect if samples
398 ..
significantly differ from one another, an
MMD is compared to its standard deviation.
If MMD>2SD, the null hypothesis
(P1=P2, where P=sample population) is
rejected at the 0·025 significance level
(Sjøvold, 1977).
Prior work suggests that as many discrete
traits as possible be used in calculating
MMD values (Sjøvold, 1977, personal com-
munication, 1993). However, they should
not be correlated, otherwise differential
weighting of underlying dimensions can
yield misleading results (Sjøvold, 1977). To
test for correlation, Kendall’s tau-b and
Spearman’s rho rank-order correlation coef-
ficient statistics were employed on the
ranked variables. The largest correlation (r)
is only 0·332 (tau-b) to 0·357 (rho)
between double shoveling UI1 and labial
curvature UI1; the total amount of variance
(r
2
)is0·11–0·13, which means only 11–13%
of the variance in the former trait is related
to that of the latter. Moreover, these particu-
lar traits were not used in the MMD analysis
(see below). The remaining trait pair corre-
lations are all much closer to zero. Thus,
there are no significant correlations among
any of the 36 traits (Carr, personal com-
munication, 1991), even though some may
be present on the same tooth (e.g., Bushman
canine UC and distal accessory ridge UC;
anterior fovea LM1 and deflecting wrinkle
LM1). The MMD statistic also requires
that rank-scale traits be divided into cat-
egories of present and absent (Sjøvold,
1977). Dichotomization was effected based
on each trait’s morphological threshold (see
Scott, 1973;Haeussler et al., 1988) accord-
ing to standard procedure (Turner, 1985b,
1987;Irish, 1993).
One of the most expeditious ways in
which to present the many distance values
among samples is via multidimensional
scaling (MDS). The procedure Alscal in
SPSS 8·0 is used in the present study.
MDS provides a spatial representation of 1
to Ndimensions consisting of a geometric
configuration of points (dental samples), as
on a map (Kruskal & Wish, 1978). Thus,
plotting samples into groups indicates
degrees of relationship.
Results
Occurrences of the 36 dental traits in the
Afalou and Taforalt samples are listed in
Table 1. The percentage of individuals in
each sample with a specific trait is presented,
along with the total number of individuals
scored. Corresponding ASU presence/
absence dichotomies follow each trait name.
To facilitate qualitative comparisons, pre-
viously published figures (Irish, 1998a,b) for
Jebel Sahaba Nubians and Capsians are
tabulated. Because of the latter’s small
sample size, several trait frequencies are
likely not representative, and should be
viewed with discretion (this issue is
addressed below). Data for the remaining
North Africans are also published elsewhere
(Irish, 1993,1998a,b), but for comparative
purposes, frequencies from a pooled group-
ing of these and other North Africans (Irish,
1993,1998a,b,c) are provided. Lastly, den-
tal data in Late Pleistocene Natufians from
Israel (Lipschultz, 1996) are listed in the
final column; they are briefly discussed later.
In prior work (Irish, 1993,1998a,b)I
described how the 13 post-Pleistocene
North African samples in the present study
show an affinity to Europeans, possessing
many traits that involve dental simplification
and mass reduction. Homogeneity of this
pattern, termed the North African Dental
Trait Complex, was reported despite vast
amounts of time (from 8000 year-old
Capsians to recent Berbers) and space (from
the Canary Islands to Egypt and Nubia)
(Irish, 1998b). It includes a comparatively
low incidence of UI1 shoveling, UC distal
accessory ridge, six-cusped LM1, LM1
deflecting wrinkle, LM2 Y-groove pattern,
and LP1 Tome’s root. In addition, there are
relatively high frequencies of four-cusped
399
Table 1 Dental trait percentages and frequencies for Afalou (AFA), Taforalt (TAF), Jebel Sahaba
(NUB), Capsian (CAP), pooled sample of North Africans (NAF),* and Natufians (NAT)†
Trait AFA TAF NUB CAP NAF NAT
Winging UI1 0·00·029·60·07·40·0
(+ =ASU 1) 0/6 0/11 8/27 0/4 34/460 0/31
Labial curvature UI1 0·014·351·933·338·4
(+=ASU 2–4) 0/3 1/7 14/27 1/3 68/177
Palatine torus 13·30·012·00·09·3
(+=ASU 2–3) 4/30 0/17 3/25 0/8 51/551
Shoveling UI1 0·040·045·80·019·53·8
(+=ASU 2–6) 0/3 2/5 11/24 0/4 30/154 6/59
Double shoveling UI1 0·00·04·30·08·612·0
(+=ASU 2–6) 0/3 0/4 1/23 0/4 15/175 12/100
Interrupt groove UI2 43·857·116·075·036·113·0
(+ =ASU+ ) 7/16 8/14 4/25 3/4 75/208 12/92
Tuberculum dentale UI2 41·77·738·975·038·895·1
(+=ASU 2–6) 5/12 1/13 7/18 3/4 73/188 58/61
Bushman canine UC 16·70·020·012·56·113·8
(+=ASU 1–3) 2/12 0/11 4/20 1/8 16/261 8/58
Distal acc. ridge UC 0·020·088·942·917·936·4
(+=ASU 2–5) 0/7 1/5 8/9 3/7 35/195 8/22
Hypocone UM2 97·0 100·094·1 100·076·791·2
(+=ASU 3–5) 32/33 20/20 32/34 9/9 342/466 134/147
Cusp 5 UM1 45·510·033·325·012·63·2
(+=ASU 2–5) 5/11 1/10 5/15 2/8 45/357 6/189
Carabelli’s UM1 41·771·450·0 100·054·781·1
(+=ASU 2–7) 5/12 5/7 7/14 4/4 181/331 73/90
Parastyle UM3 3·10·00·00·01·20·8
(+=ASU 1–5) 1/32 0/18 0/37 0/8 4/332 1/133
Enamel extension UM1 0·00·058·30·06·80·0
(+=ASU 1–3) 0/31 0/21 21/36 0/12 34/503 0/156
Root No. UP1 55·662·572·736·457·166·7
(+ =ASU 2+) 15/27 10/16 24/33 4/11 267/468 22/33
Root No. UM2 87·562·573·085·778·693·5
(+ =ASU 3+) 14/16 10/16 27/37 6/7 294/374 43/46
Peg-reduced UI2 0·00·02·90·01·8
(+ =ASU P or R) 0/33 0/27 1/34 0/8 5/275
Odontome P1-2 20·00·00·00·00·20·6
(+ =ASU+ ) 2/10 0/10 0/11 0/11 1/144 1/161
Congenital absence UM3 2·94·50·018·215·2
(+=ASU) 1/35 1/22 0/45 2/11 83/545
Midline diastema UI1 0·00·023·10·06·1
(+=d0·5 mm) 0/6 0/13 6/26 0/4 25/413
Lingual cusp LP2 89·535·793·383·372·660·2
(+=ASU 2–9) 17/19 5/14 14/15 10/12 196/270 59/98
Anterior fovea LM1 0·08·369·250·037·9
(+=ASU 2–4) 0/12 1/12 9/13 5/10 75/198
Mandibular torus 0·00·00·00·01·0
(+=ASU 2–3) 0/28 0/25 0/47 0/17 5/493
Groove pattern LM2 79·334·862·533·330·630·5
(+ =ASU Y) 23/29 8/23 20/32 5/15 123/402 47/154
Rocker jaw 13·829·20·020·019·3
(+=ASU 1–2) 4/29 7/24 0/45 3/15 93/481
Cusp No. LM1 11·113·331·320·07·719·2
(+ =ASU 6+) 2/18 2/15 10/32 3/15 27/352 30/156
Cusp No. LM2 76·910·594·631·333·640·9
(+ =ASU 5+) 20/26 2/19 35/37 5/16 128/381 72/176
400 ..
LM2 and M3 agenesis, as well as UM1
Carabelli’s trait and two-rooted LC. Rocker
jaw,
1
a common European feature (Turner
& Markowitz, 1990), is also seen. West
Asians are reported to possess many similar
traits (Roler, 1992;Lipschultz, 1996),
although comprehensive study of these
populations has yet to be undertaken. Any
North African deviations way from this
simple dental pattern are in the direction
of complex, sub-Saharan traits (e.g., UI2
tuberculum dentale, Bushman canine, two-
rooted UP1, three-rooted UM2), suggesting
some admixture with these peoples (Irish,
1993,1997). These findings agree with
genetic-based results that link North
Africans to Europeans and West Asians,
but record many sub-Saharan influenced
markers (e.g., Mourant, 1983;Hiernaux,
1975;Nurse et al., 1985;Sanchez-
Mazas et al., 1986;Excoffier et al., 1987;
Roychoudhury & Nei, 1988;Arnaiz-Villena
et al., 1997).
Iberomaurusian dental trait frequencies
suggest that the time depth for the North
African pattern may be pushed back farther
than formerly envisioned (Irish, 1998b).
Taforalt teeth possess simple morphology,
and include such diagnostic traits as the UI2
interruption groove, LM2, +-groove pat-
tern, four-cusped LM2, M3 agenesis (or
reduction), and rocker jaw. The Afalou
dentitions show a similar pattern, albeit
with a trend toward greater morphological
complexity. Such traits as five-cusped UM1,
LM2 Y-groove pattern, five-cusped LM2,
and LP1 Tome’s root are present in
markedly higher frequencies (see Table 1).
Dentitions of Late Pleistocene Jebel
Sahaba Nubians have extremely high fre-
quencies of complex, mass-additive (and
1
Although the ASU System’s rocker jaw trait is most
often associated with unrelated Polynesians (Turner &
Scott, 1977), its second highest worldwide incidence is
reported among Europeans and other nearby peoples
(e.g., Turner & Markowitz, 1990). It thus serves, along
with other oral discrete traits, as an excellent discrimi-
nator between European-like peoples and Asians and
sub-Saharan Africans.
Table 1 Continued
Trait AFA TAF NUB CAP NAF NAT
Deflecting wrinkle LM1 0·00·030·822·28·23·4
(+=ASU 2–3) 0/14 0/11 4/13 2/9 22/267 4/119
C1-C2 crest LM1 0·00·00·00·03·30·0
(+ =ASU+ ) 0/6 0/8 0/16 0/8 9/276 0/119
Protostylid LM1 15·030·829·246·232·514·4
(+=ASU 1–6) 3/20 4/13 7/24 6/13 114/351 21/146
Cusp 7 LM1 14·83·89·718·85·12·9
(+=ASU 2–4) 4/27 1/26 3/31 3/16 21/414 5/170
Tome’s root LP1 21·10·052·40·08·620·0
(+=ASU 3–5) 4/19 0/18 11/21 0/14 32/372 2/10
Root No. LC 0·00·00·00·02·34·0
(+ =ASU 2+) 0/19 0/19 0/17 0/11 8/347 1/25
Root No. LM1 0·00·013·06·21·20·0
(+ =ASU 3+) 0/12 0/22 6/46 1/16 4/337 0/23
Root No. LM2 100·075·083·785·788·3100·0
(+ =ASU 2+) 8/8 12/16 36/43 12/14 294/333 33/33
Torsomolar angle LM3 0·04·27·727·321·6
(+ =ASU+ ) 0/23 1/24 3/39 3/11 74/343
*Nubian, Capsian, and pooled North African comparative data from Irish (1993,1998a,b) (see text for details).
†Natufian comparative data from Lipschultz (1996).
401
other) traits, including UI1 labial curvature,
UI1 shoveling, Bushman Canine, UC distal
accessory ridge, midline diastema, six-
cusped LM1, LM2 Y-5, and LP1 Tome’s
root. Furthermore, they exhibit low fre-
quencies of typical North African features.
This trait combination is ubiquitous in sub-
Saharan Africans (Irish & Turner, 1990;
Irish, 1993,1997,1998a,b, for details).
These qualitative comparisons are sup-
ported by the MMD results.
2
Unfortu-
nately, due to the Iberomaurusian UI1
evulsion, five corresponding traits (winging
labial curvature, shoveling, double
shoveling, and midline diastema) were
excluded from analysis because of small
sample sizes (i.e., one or both samples <8).
Distal accessory ridge UC and C1–C2 crest
LM1 were dropped for the same reason.
The remaining 29 traits were used to deter-
mine biological distances. Analysis of the
small Capsian sample was also ameliorated
by these adjustments. MMD distances
among the samples are illustrated via
interval-level, two-dimensional MDS in
Figure 2. This measurement level was
decided to be most appropriate, because the
large number of traits causes the matrix of
distance values to approximate continuous
data (Carr, personal communication, 1991).
The easily interpretable two-dimensional
configuration is used, rather than three-, due
to minimal improvement in stress and r
2
values at the higher dimension.
Taforalt (far left) is associated with the 13
closely clustered, post-Pleistocene samples.
Within this cluster, mingling of northwest
and northeast Africans reflects the dental
homogeniety previously mentioned. MMD
values among the 13 samples range between
0·00 and 0·08; most are not significantly
different. MMDs between Taforalt and
these samples vary from 0·02 to 0·14;
seven values are not significant, includ-
ing Capsians (0·06) and Shawia Berbers
(0·04). Nonsignificant MMDs also occur
between Taforalt and Carthaginians (0·02),
Soleb Nubians (0·02), Christian Nubians
(0·04), Kharga Egyptians (0·01), and Lisht
Egyptians (0·05). Jebel Sahaba (far
right) is significantly divergent from all
others (MMD range=0·22–0·47). Although
2
A table of distance values among all samples is
available upon request from the author.
Figure 2. Multidimensional scaling of distance values among 16 North African samples.
402 ..
simplistic, the x-axis can be envisioned as a
line representing increasing dental complex-
ity. Therefore, Taforalt has the most mass-
reduced, while Jebel Sahaba has the most
mass-additive features. Finally, Afalou (bot-
tom) shows obvious separation from the
rest, although, compared to Jebel Sahaba, it
could possibly be seen as an outlier of the
main cluster using a neighborhood approach
to MDS cluster interpretation (see
Guttman, 1954;Kruskal & Wish, 1978).
MMDs between Afalou and post-
Pleistocene samples are 0·08 to 0·27,
whereas the corresponding value between
Afalou and Taforalt is 0·17; all are signifi-
cant. Analogous results (Irish, 1999) were
obtained using another distance measure
[i.e., Konigsberg’s modified Mahalanobis
(Konigsberg, 1990;Ishida & Dodo, 1997)],
suggesting the affinities are actual, not an
artefact of a particular statistical method.
Discussion and conclusions
How do these numerically derived findings
relate to the questions presented in the
introduction? First, the idea that Iberomau-
rusians are unrelated to subsequent North
Africans is not supported. With Taforalt, at
least, these is no conspicuous divergence to
suggest they were a genetic dead-end.
Second, Iberomaurusians are wholly unlike
Late Pleistocene Nubians, despite pur-
ported similarities in cultural manifestations
and cranial robusticity. Indeed, Taforalt and
Jebel Sahaba appear to be at opposite ends
of a dental morphological spectrum.
With respect to population continuity,
both Iberomaurusian samples show some
degree of affiliation with all later North
Africans, as suggested by the sharing of
many morphologically simple features found
in the North African Dental Trait Complex.
Taforalt exhibits the closest affiliation, based
on its proximity to the post-Pleisocene clus-
ter. Within the Maghreb, Taforalt is most
akin to the Shawia Berbers and Capsians,
although the small Capsian sample
requires that these results be interpreted
with caution; Afalou shows slight similitude
to Canary Island Guanches. Together, these
affinities may be indicative of regional
population continuity, which supports sev-
eral findings by Close (1986), Lubell and
co-workers (Lubell et al., 1984;Sheppard
& Lubell, 1990;Lubell, 2000), and per-
haps those hypothesizing an Iberomau-
rusian/Guanche connection (Vallois, 1969;
Ferembach, 1985). However, there is no
evidence for a close affinity between Afalou
and Capsians, or with most other samples.
Recent cranial analyses (Groves & Thorne,
1999) support these findings. Thus,
conversely, several conclusions by Balout
(1955),Chamla (1973,1975,1978),
Camps (1974), and Hiernaux (1975) are
sustained. To further confuse matters,
Taforalt is allied with West Asian-derived
Carthaginians, and recent Egyptians and
Nubians. If not perceived as an indicator of
West Asian (e.g., Vallois, 1969) or Nile
Valley (Lubell et al., 1984)influence, these
affinities may simply represent an overall
correspondence with the mass-reduced den-
tal pattern prevalent in greater post-
Pleistocene North Africa. The Capsian
sample is also linked with some Nubians
(Figure 2). Again, if this is not seen as sup-
port for a Capsian origin in the east (Briggs,
1954,1955;Camps, 1974;Chamla, 1978;
Ferembach, 1962,1985), it may represent a
fortuitous relationship due to sample size
and heterogeneity, or perhaps is sympto-
matic of pervasive post-Pleistocene dental
homogeneity.
There are more definitive answers regard-
ing the question of homogeneity between
Late Pleistocene Iberomaurusians and
Nubians. Extreme divergence between the
two suggests they are not closely related.
Whereas Afalou and, particularly, Taforalt
dentitions are characterized by dental mor-
phological reduction, the Nubians exhibit a
mass-additive dental pattern, like that in
403
sub-Saharan peoples. The latter possess a
suite of 11 mass-additive traits that I termed
the Sub-Saharan African Dental Complex
(Irish, 1997,1998a). These findings bolster
previous dental (Irish & Turner, 1990,
1992;Irish, 1993,1997,1998a,b,c,1999),
cranial (Franciscus,1995, personal com-
munication, 1996; Groves & Thorne,
1999), and postcranial (Holliday, 1995)
results.
Thus, evidence for a common Mechta-
Afalou population in both the Maghreb and
Nubia (Anderson, 1968;Clark, 1970;
Greene & Armelagos, 1972;Ferembach,
1985;Dutour, 1995;Lahr & Arensburg,
1995) is not supported. Calls for similarity
based on such shared cranial features as
prominent brows, projecting zygomatic
arches, gonial eversion, alveolar prognath-
ism, and complex teeth, among others
(Anderson, 1968;Greene, 1972;Greene &
Armelagos, 1972), may be ill founded.
Except for several Afalou traits, it was dem-
onstrated that Iberomaurusians do not pos-
ses complex teeth. Moreover, even a casual
inspection of crania in the three samples (see
Figure 3) reveals that many characteristic
Nubian traits, including, for example,
alveolar prognathism, are uncommon or
absent in Iberomaurusians (see Groves &
Thorne, 1999 for more detailed comparison
of traits).
Lastly, an additional, though not com-
pletely unexpected, finding entails the sig-
nificant divergence between the two Ibero-
maurusian samples. Besides shared cranial
traits, which can result from common
environmental factors (e.g., Smith, 1979),
the Taforalt and Afalou-Bou-Rhummel
samples differ in a number of dental trait
frequencies. As Ferembach (1962) notes,
such morphological variations should not be
too surprising, considering the two samples
were separated by 700 km and up to
3000years. Intra-Iberomaurusian diver-
sity has been observed in terms of racial
typologies, including Briggs’(1955) A–D
cranial types, Chamla’s (1978:393) desig-
nation of ‘‘Typique’’ and ‘‘Gracilise´’’, and
Camps’(1974:162) ‘‘Mechta-Afalou clas-
sique’’ and ‘‘Mechtoı¨des e´volue´s’’. How-
ever, whether a result of sub-Saharan gene
flow (see Lahr & Arensburg, 1995;Groves
& Thorne, 1999) into Afalou, other genetic
factors (Chamla, 1973), or even dietary and
behavioral variation (Smith, 1979), the
diversity provides additional evidence for
Late Pleistocene population heterogeneity in
the region. Therefore, it may be unwise to
pool Taforalt and Afalou into a single Ibero-
maurusian sample, as has often been the
case in previous studies (e.g., Chamla,
1980;Bermudez de Castro, 1991).
Future work to address these and related
issues will entail the collection of more den-
tal data, with an emphasis on the pivotal
Capsian sample, and Late Pleistocene and
subsequent West Asians, among others. To
briefly illustrate the importance of incorpor-
ating, for example, West Asians, this study
concludes with a 22-trait comparison
between the North Africans and a sample of
Late Pleistocene (ca. 12,800–10,200 BP)
Natufians (NAT) from Israel (Figure 4).
Data from these latter individuals are in
Lipschultz (1996); they were also recorded
using the ASU System. Twenty-two rather
than 29 traits (see Table 1) are compared
using the MMD statistic because the other
seven are not listed in his thesis.
Although interobserver error cannot be
ruled out as a factor, Natufians (top of
Figure 4) are significantly divergent from
Iberomaurusians and other North Africans
(MMD range=0·10–0·43). Despite contem-
poraneity, they differ most from Afalou
(0·27), Taforalt (0·27), and Jebel Sahaba
(0·43). These findings support conclusions
by Ferembach (1962),Camps (1974),
Hershkovitz et al. (1995),Lahr & Arensburg
(1995), and others (see Dutour, 1995)
based on skeletal metric and nonmetric
data. It also suggests Late Pleistocene
population heterogeneity extended beyond
404 ..
Figure 3. Lateral views of three Late Pleistocene male crania showing alveolar prognathism in Jebel
Sahaba 117-10 (top), but not in Taforalt XI-C1 (middle) or Afalou 3 (bottom).
405
North Africa into the greater circum-
Mediterranean area, as suggested by Smith
(1979). Lastly, the Natufian lack of affinity
to Caspians (0·16) is contra Ferembach’s
(1962) mention of a possible ancestor–
descendent relationship. Whatever the case,
clearly much remains to be worked out in
the area, and comprehensive analyses of
extra-African samples are likely to be a key
in learning the ultimate origins and affinities
of Late Pleistocene through Recent North
African peoples.
Summary
With reference to a literature review of
North African population history, 36 non-
metric dental traits in two Late Pleisto-
cene Iberomaurusian samples from
Taforalt, Morocco, and Afalou-Bou-
Rhummel, Algeria, were contrasted with one
another, and with those in contemporaneous
Jebel Sahaba Nubians, post-Pleistocene
Maghreb Capsians, and 12 younger north-
west and northeast African samples.
Comparisons of trait frequencies, and results
from the Mean Measure of Divergence dis-
tance statistic using 29 of the 36 traits, facili-
tated estimates of biological affinities among
samples. Taforalt Iberomaurusians are simi-
lar to many post-Pleistocene North Africans,
including such Maghreb groups as Capsians
and Shawia Berbers. Compared to Taforalt,
Afalou Iberomaurusians are divergent from
all samples, though they show a distant
affinity to Canary Island Gaunches. Both
Iberomaurusian samples, as well as post-
Pleistocene North Africans, are extremely
divergent from Late Pleistocene Nubians.
The latter are more akin to sub-Saharan
Africans. All told, these numerically-derived
findings provide some level of support for
genetic continuity between, at least, Taforalt
Iberomaurusians and later populations in
the Maghreb and greater North Africa since
the terminal Pleistocene. However, popu-
lation heterogeneity was apparently the rule
before that time.
Acknowledgements
An earlier version of this paper was pre-
sented at the symposium ‘‘Antropologia
dentale delle popolazioni pre-protostoriche
Figure 4. Multidimensional scaling of distance values among 16 North African samples and Natufians.
406 ..
del Mediterraneo’’, chaired by Alfredo
Coppa, Universita´di Roma ‘‘La Sapienza’’,
for the XIII Congresso degli antropologi
italiani, Roma e Sabaudia, October, 1999.
Funding to attend the meeting was coordi-
nated through Prof. Coppa. David Lubell
from the University of Alberta kindly
provided a pre-publication copy of his
Late Pleistocene–Early Holocene Maghreb
article, and other useful materials. I also
thank Dr Lubell, Mary Jackes (University of
Alberta), Terry Harrison (Joint JHE Editor),
and the JHE reviewers for providing
constructive criticism and comments on an
earlier version of this paper. Lastly, I am
grateful to the many individuals at the insti-
tutions where I collected data, including:
Christy Turner, Charles Merbs and Donald
Morris from Arizona State University
(ASU), Tempe, AZ; Guy Gibbon and the
late Elden Johnson from the University of
Minnesota, Minneapolis, MN; Douglas
Ubelaker and David Hunt from the
National Museum of Natural History,
Washington, DC; Ian Tattersall, Jaymie
Brauer, and Gary Sawyer from the
American Museum of Natural History, New
York; Andre Langaney, Frances Roville-
Sausse, and Miya Awazu Periera da Silva
from the Muse´edel’Homme, Paris; Fred
Wendorf and Sue Linder-Linsley from
Southern Methodist University, Dallas, TX;
and Henry de Lumley and Dominique
Grimaud-Herve, Institut de Pale´onologie
Humaine, Paris. The comparative data
were collected via support from the
National Science Foundation (BNS-
9013942), ASU Research Development
Program, and American Museum of Natural
History.
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