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 anity 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 anities. 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, 393410
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
latters 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
supercial?
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 20,000 years 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 997757720, U.S.A. Tel.:
(907) 474-6755; E-mail: jdi@uaf.edu
00472484/00/100393+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 latters 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
akes, 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
eastperhaps from as far aeld 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 inuence, it prob-
ably came from the Nile Valley (i.e., Late
Pleistocene Nubians) rather than West Asia.
Concerning the question of anity to
contemporaneous peoples, previous studies
show that Late Pleistocene Nubians share a
number of similarities with Iberomauru-
sians; the formers 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 . b. inisn
(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 Castros (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 diers
noticeably from that of the Iberomauru-
sians, and describes several physical dier-
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 anity; 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 ndings are largely
supported by Groves & Thornes (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 Paleontologie 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
Hachis (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 Paleontologie
Humaine. Although background records are
395 incno:acncsiaN arriNi1ics
incomplete, this small, heterogenous sample
comes from both Algerian and Tunisian
sites [including Mechta el-Arbi, Mechta-
Chateaudun, A 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, A 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
Musee de lHomme. Greenberg (1966)
characterizes Berbers as speaking one of
several dialects (e.g., Shawia) of a native
Berber language, which reects an inuence
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 Musee de lHomme. 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
Musee de lHomme, 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 ow from
Arabs and Carthaginians (Schwidetsky,
Figure 1. Origin locations of the 16 North African dental samples.
396 . b. inisn
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. 14,50012,500 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 ani-
ties at a regional level. Three samples com-
prise 12th Dynasty through Byzantine
Egyptians (40001400 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 (35753380 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 (2100600 BP) from Semna
(Zabkar & Zabkar, 1982). The Pharonic
sample was recorded at the Musee de
lHomme; 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 Musee de
lHomme 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 Musee de
lHomme. Arabs rst 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 incno:acncsiaN arriNi1ics
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 anity 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 individuals
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.
Smiths Mean Measure of Divergence
(MMD) distance statistic, using the
Freeman and Tukey angular transformation
correction (Berry & Berry, 1967; Sjvold,
1973; Green & Suchey, 1976) for small
sample sizes, and low (c005) or high
(d095) trait frequencies (Sjvold, 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 anity, 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 . b. inisn
signicantly dier 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 0025 signicance level
(Sjvold, 1977).
Prior work suggests that as many discrete
traits as possible be used in calculating
MMD values (Sjvold, 1977, personal com-
munication, 1993). However, they should
not be correlated, otherwise dierential
weighting of underlying dimensions can
yield misleading results (Sjvold, 1977). To
test for correlation, Kendalls tau-b and
Spearmans rho rank-order correlation coef-
cient statistics were employed on the
ranked variables. The largest correlation (r)
is only 0332 (tau-b) to 0357 (rho)
between double shoveling UI1 and labial
curvature UI1; the total amount of variance
(r
2
) is 011013, which means only 1113%
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 signicant 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 deecting wrinkle
LM1). The MMD statistic also requires
that rank-scale traits be divided into cat-
egories of present and absent (Sjvold,
1977). Dichotomization was eected based
on each traits 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 80 is used in the present study.
MDS provides a spatial representation of 1
to N dimensions consisting of a geometric
conguration 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 specic 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 gures (Irish, 1998a,b) for
Jebel Sahaba Nubians and Capsians are
tabulated. Because of the latters 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 Natuans from
Israel (Lipschultz, 1996) are listed in the
nal column; they are briey 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 anity to Europeans, possessing
many traits that involve dental simplication
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
deecting wrinkle, LM2 Y-groove pattern,
and LP1 Tomes root. In addition, there are
relatively high frequencies of four-cusped
399 incno:acncsiaN arriNi1ics
Table 1 Dental trait percentages and frequencies for Afalou (AFA), Taforalt (TAF), Jebel Sahaba
(NUB), Capsian (CAP), pooled sample of North Africans (NAF),* and Natuans (NAT)
Trait AFA TAF NUB CAP NAF NAT
Winging UI1 00 00 296 00 74 00
(+=ASU 1) 0/6 0/11 8/27 0/4 34/460 0/31
Labial curvature UI1 00 143 519 333 384
(+=ASU 24) 0/3 1/7 14/27 1/3 68/177
Palatine torus 133 00 120 00 93
(+=ASU 23) 4/30 0/17 3/25 0/8 51/551
Shoveling UI1 00 400 458 00 195 38
(+=ASU 26) 0/3 2/5 11/24 0/4 30/154 6/59
Double shoveling UI1 00 00 43 00 86 120
(+=ASU 26) 0/3 0/4 1/23 0/4 15/175 12/100
Interrupt groove UI2 438 571 160 750 361 130
(+=ASU+) 7/16 8/14 4/25 3/4 75/208 12/92
Tuberculum dentale UI2 417 77 389 750 388 951
(+=ASU 26) 5/12 1/13 7/18 3/4 73/188 58/61
Bushman canine UC 167 00 200 125 61 138
(+=ASU 13) 2/12 0/11 4/20 1/8 16/261 8/58
Distal acc. ridge UC 00 200 889 429 179 364
(+=ASU 25) 0/7 1/5 8/9 3/7 35/195 8/22
Hypocone UM2 970 1000 941 1000 767 912
(+=ASU 35) 32/33 20/20 32/34 9/9 342/466 134/147
Cusp 5 UM1 455 100 333 250 126 32
(+=ASU 25) 5/11 1/10 5/15 2/8 45/357 6/189
Carabellis UM1 417 714 500 1000 547 811
(+=ASU 27) 5/12 5/7 7/14 4/4 181/331 73/90
Parastyle UM3 31 00 00 00 12 08
(+=ASU 15) 1/32 0/18 0/37 0/8 4/332 1/133
Enamel extension UM1 00 00 583 00 68 00
(+=ASU 13) 0/31 0/21 21/36 0/12 34/503 0/156
Root No. UP1 556 625 727 364 571 667
(+=ASU 2+) 15/27 10/16 24/33 4/11 267/468 22/33
Root No. UM2 875 625 730 857 786 935
(+=ASU 3+) 14/16 10/16 27/37 6/7 294/374 43/46
Peg-reduced UI2 00 00 29 00 18
(+=ASU P or R) 0/33 0/27 1/34 0/8 5/275
Odontome P1-2 200 00 00 00 02 06
(+=ASU+) 2/10 0/10 0/11 0/11 1/144 1/161
Congenital absence UM3 29 45 00 182 152
(+=ASU) 1/35 1/22 0/45 2/11 83/545
Midline diastema UI1 00 00 231 00 61
(+=d05 mm) 0/6 0/13 6/26 0/4 25/413
Lingual cusp LP2 895 357 933 833 726 602
(+=ASU 29) 17/19 5/14 14/15 10/12 196/270 59/98
Anterior fovea LM1 00 83 692 500 379
(+=ASU 24) 0/12 1/12 9/13 5/10 75/198
Mandibular torus 00 00 00 00 10
(+=ASU 23) 0/28 0/25 0/47 0/17 5/493
Groove pattern LM2 793 348 625 333 306 305
(+=ASU Y) 23/29 8/23 20/32 5/15 123/402 47/154
Rocker jaw 138 292 00 200 193
(+=ASU 12) 4/29 7/24 0/45 3/15 93/481
Cusp No. LM1 111 133 313 200 77 192
(+=ASU 6+) 2/18 2/15 10/32 3/15 27/352 30/156
Cusp No. LM2 769 105 946 313 336 409
(+=ASU 5+) 20/26 2/19 35/37 5/16 128/381 72/176
400 . b. inisn
LM2 and M3 agenesis, as well as UM1
Carabellis 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 ndings agree with
genetic-based results that link North
Africans to Europeans and West Asians,
but record many sub-Saharan inuenced
markers (e.g., Mourant, 1983; Hiernaux,
1975; Nurse et al., 1985; Sanchez-
Mazas et al., 1986; Excoer 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 ve-cusped UM1,
LM2 Y-groove pattern, ve-cusped LM2,
and LP1 Tomes 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 Systems 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
Deecting wrinkle LM1 00 00 308 222 82 34
(+=ASU 23) 0/14 0/11 4/13 2/9 22/267 4/119
C1-C2 crest LM1 00 00 00 00 33 00
(+=ASU+) 0/6 0/8 0/16 0/8 9/276 0/119
Protostylid LM1 150 308 292 462 325 144
(+=ASU 16) 3/20 4/13 7/24 6/13 114/351 21/146
Cusp 7 LM1 148 38 97 188 51 29
(+=ASU 24) 4/27 1/26 3/31 3/16 21/414 5/170
Tomes root LP1 211 00 524 00 86 200
(+=ASU 35) 4/19 0/18 11/21 0/14 32/372 2/10
Root No. LC 00 00 00 00 23 40
(+=ASU 2+) 0/19 0/19 0/17 0/11 8/347 1/25
Root No. LM1 00 00 130 62 12 00
(+=ASU 3+) 0/12 0/22 6/46 1/16 4/337 0/23
Root No. LM2 1000 750 837 857 883 1000
(+=ASU 2+) 8/8 12/16 36/43 12/14 294/333 33/33
Torsomolar angle LM3 00 42 77 273 216
(+=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).
Natuan comparative data from Lipschultz (1996).
401 incno:acncsiaN arriNi1ics
other) traits, including UI1 labial curvature,
UI1 shoveling, Bushman Canine, UC distal
accessory ridge, midline diastema, six-
cusped LM1, LM2 Y-5, and LP1 Tomes
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, ve 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 C1C2 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
conguration 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 reects the dental
homogeniety previously mentioned. MMD
values among the 13 samples range between
000 and 008; most are not signicantly
dierent. MMDs between Taforalt and
these samples vary from 002 to 014;
seven values are not signicant, includ-
ing Capsians (006) and Shawia Berbers
(004). Nonsignicant MMDs also occur
between Taforalt and Carthaginians (002),
Soleb Nubians (002), Christian Nubians
(004), Kharga Egyptians (001), and Lisht
Egyptians (005). Jebel Sahaba (far
right) is signicantly divergent from all
others (MMD range=022047). 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 . b. inisn
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 008 to 027,
whereas the corresponding value between
Afalou and Taforalt is 017; all are signi-
cant. Analogous results (Irish, 1999) were
obtained using another distance measure
[i.e., Konigsbergs modied Mahalanobis
(Konigsberg, 1990; Ishida & Dodo, 1997)],
suggesting the anities are actual, not an
artefact of a particular statistical method.
Discussion and conclusions
How do these numerically derived ndings
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 aliation 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 aliation, 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
anities may be indicative of regional
population continuity, which supports sev-
eral ndings 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 anity between Afalou
and Capsians, or with most other samples.
Recent cranial analyses (Groves & Thorne,
1999) support these ndings. 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) inuence, these
afnities 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 denitive 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 incno:acncsiaN arriNi1ics
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 ndings 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, nding entails the sig-
nicant 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 dier 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) AD
cranial types, Chamlas (1978:393) desig-
nation of Typique and Gracilise, and
Camps (1974:162) Mechta-Afalou clas-
sique and Mechto des evolues. How-
ever, whether a result of sub-Saharan gene
ow (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
briey 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. 12,80010,200 BP)
Natuans (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, Natuans (top of
Figure 4) are signicantly divergent from
Iberomaurusians and other North Africans
(MMD range=010043). Despite contem-
poraneity, they dier most from Afalou
(027), Taforalt (027), and Jebel Sahaba
(043). These ndings 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 . b. inisn
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 incno:acncsiaN arriNi1ics
North Africa into the greater circum-
Mediterranean area, as suggested by Smith
(1979). Lastly, the Natuan lack of anity
to Caspians (016) is contra Ferembachs
(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 anities
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 anities 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
anity 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
ndings 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 Natuans.
406 . b. inisn
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 PleistoceneEarly 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 Musee de lHomme, Paris; Fred
Wendorf and Sue Linder-Linsley from
Southern Methodist University, Dallas, TX;
and Henry de Lumley and Dominique
Grimaud-Herve, Institut de Paleonologie
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.
References
Anderson, J. E. (1968). Late Paleolithic skeletal
remains from Nubia. In (F. Wendorf, Ed.) The
Prehistory of Nubia, Vol. 2, pp. 9961040. Dallas:
Fort Burgwin Research Center.
Angel, J. L. & Kelley, J. O. (1986). Description and
comparison of the skeleton. In (A. Close, Ed.) The
Prehistory of Wadi Kubbaniya. Volume 1: The Wadi
Kubbaniya Skeleton: A Late Paleolithic Burial
from Southern Egypt, pp. 5370. Dallas: Southern
Methodist University Press.
Arnaiz-Villena, A., Martinez-Laso, J., Gomez-Casado,
E., Diaz-Campos, N., Santos, P., Martinho, A. &
Breda-Coimbra, H. (1997). Relatedness among
Basques, Portuguese, Spaniards, and Algerians
studied by HLA allelic frequencies and haplotypes.
Immunogenetics 47, 3743.
Baines, J. & Malek, J. (1982). Atlas of Ancient Egypt.
New York: Facts On File.
Balout, L. (1955). Prehistoire de lAfrique du Nord. Paris:
Arts et Metiers Graphiques.
Balout, L. & Briggs, L. C. (1949). Debris humains de
lescargotie`re de Gambetta. Bull. Soc. dHist. Nat.
lAfrique Nord 40, 125134.
Bermudez de Castro, J. M. (1989). The Carabelli trait
in human prehistoric populations of the Canary
Islands. Hum. Biol. 61, 117131.
Bermudez de Castro, J. M. (1991). La denture
de la population Mesolithique dAfrique du
Nord et lhypothe`se Afro-Europeen Sapiens.
LAnthropologie 95, 201218.
Berry, A. C. & Berry, R. J. (1967). Epigenetic variation
in the human cranium. J. Anat. 101, 361379.
Briggs, L. C. (1954). Deux te`tes osseuses de la collec-
tion Debruge, le crane type de Mechta-el-Arbi et
le crane A de la grotte des Hye`nes. Libyca 2,
121149.
Briggs, L. C. (1955). The Stone Age Races of Northwest
Africa. Bulletin of the American School of Prehistoric
Research, No. 18. Cambridge: Peabody Museum.
Bynon, J. (1970). The contribution of linguistics to
history in the eld of Berber studies. In (D. Dalby,
Ed.) Language and History in Africa, pp. 6477. New
York: Africana Publishing.
Caillard, P. (1978). Dimensions dentaires et usure
dentaire proximale de lHomme dAfalou-Bou-
Rhummel. Bull. Mem. Soc. Anthropol. Paris 5, 261
274.
Camps, G. (1974). Les Civilisations Prehistoriques de
lAfrique du Nord et du Sahara. Paris: Doin.
Camps-Fabrer, H. (1975). Un Gisement Capsien de
Facie`s Setien Medjez II, El Eulma, (Algerie). Paris:
CNRS.
Carlson, D. S. & Van Gerven, D. P. (1977). Mastica-
tory function and post-Pleistocene evolution in
Nubia. Am. J. phys. Anthrop. 46, 495506.
Chabeuf, M. (1975). Eutude anthropologique de
Medjez II. In (H. Camps-Fabrer, Ed.) Un Gisement
Capsien de Facie`s Setien Medjez II, El Eulma,
(Algerie), pp. 331371. Paris: CNRS.
Chamla, M. C. (1973). Etude anthropologique de
lHomme capsien de lA n Dokkara (Algerie orien-
tale). Libyca 21, 953.
Chamla, M. C. (1975). La diversite des types humains
dans les gisements capsiens. In (H. Camps-Fabrer,
Ed.) Un Gisement Capsien de Facie`s Setien Medjez II,
El Eulma, (Algerie), pp. 373376. Paris: CNRS.
407 incno:acncsiaN arriNi1ics
Chamla, M. C. (1978). Le peuplement de lAfrique du
Nord de lEpipaleolithique a` lepoque actuelle.
LAnthropologie 82, 385430.
Chamla, M. C. (1980). Eutude des variations metriques
des couronnes dentaires des Nord-fricains, de lEupi-
paleolithique a` lepoque actuelle. LAnthropologie 84,
254271.
Charles-Picard, G. & Picard, C. (1968). The Life and
Death of Carthage. New York: Taplinger.
Clark, J. D. (1970). The Prehistory of Africa. New York:
Praeger.
Clark, J. D. (1989). The origins and spread of modern
humans: a broad perspective on the African evidence.
In (P. Mellars & C. Stringer, Eds) The Human
Revolution: Behavioural and Biological Perspectives
on the Origins of Modern Humans, pp. 565588.
Edinburgh: Edinburgh University Press.
Close, A. (1986). The place of the Haua Fteah in the
Late Palaeolithic of North Africa. In (G. N. Bailey &
P. Callow, Eds) Stone Age Prehistory: Studies
in Memory of Charles McBurney, pp. 169180.
Cambridge: Cambridge University Press.
Close, A. E. (1977). The Identication of Style in Lithic
Artefacts from North-East Africa. Cairo: Memoires
delInstitut dEgypte, No. 61.
Cluzel, C. (1973). Euvolution de la gracilisation dentaire
dans les anciennes populations du Maghreb. Libyca
21, 5561.
Curto, S. (1972). Archaeological outline of Egypt from
the Paleolithic to the modern Arab state. J. hum.
Evol. 1, 141146.
Dahlberg, A. A. (1963). Analysis of the American
Indian dentition. In (D. Brothwell, Ed.) Dental
Anthropology, pp. 149158. New York: Pergamon
Press.
Davidson, B. (1974). Africa in History. New York:
Macmillan.
Dutour, O. (1995). Le peuplement modern dAfrique
septentrionale et ses relations avec celui du Proche-
Orient. Paleorient 21, 97109.
Elliot Smith, G. & Wood-Jones, F. (1910). The
Archaeological Survey of Nubia: Report for 19071908,
Volume II, Report on Human Remains. Cairo: National
Printing Department.
Excoer, L., Pellegrini, B., Sanchez-Mazas, A., Simon,
C. & Langaney, A. (1987). Genetics and history of
sub-Saharan Africa. Yearb. phys. Anthrop. 30, 151
194.
Ferembach, D. (1962). La Necropole Eupipaleolithique de
Taforalt (Maroc Oriental). Etude des Squelettes
Humains. Paris: CNRS.
Ferembach, D. (1985). On the origin of the Iberomau-
rusians. A new hypothesis. J. hum. Evol. 14, 393397.
Franciscus, R. (1995). Nasal morphology in western
Old World Later Pleistocene hominids and the
origins of modern humans. Ph.D. Dissertation,
University of New Mexico, Albuquerque.
Green, R. & Suchey, J. (1976). The use of inverse sine
transformation in the analysis of non-metrical data.
Am. J. phys. Anthrop. 45, 6168.
Greenberg, J. H. (1966). The Languages of Africa.
Bloomington: Indiana University.
Greene, D. L. (1967). Dentition of Meroitic, X-Group,
and Christian Populations From Wadi Halfa, Sudan.
Anthropol. Paper 85, Nubian Ser. 1. Salt Lake City:
University of Utah Press.
Greene, D. L. (1972). Dental anthropology of early
Egypt and Nubia. J. hum. Evol. 1, 315324.
Greene, D. L. & Armelagos, G. (1972). The Wadi Halfa
Mesolithic Population. Res. Report 11. Dept. Anthro-
pology. Amherst: University of Massachusetts.
Groves, C. P. & Thorne, A. (1999). The terminal
Pleistocene and early Holocene populations of
northern Africa. Homo 50, 249262.
Guttman, L. (1954). Some necessary conditions for
common-factor analysis. Psychometrika 19, 149161.
Hachi, S. (1996). LIberomaurusien, decouverte des
fouilles dAfalou (Bedja a, Algerie). LAnthropologie
100, 5576.
Hachi, S. (1997). Resultats des fouilles recentes
dAfalou Bou Rmel (Bedja a, Algerie). In (J. M.
Fullola & N. Soler, Eds) El mon mediterrani despre del
Pleniglacial (18,00012,000 BP). Se`rie Monogra`ca,
17, pp. 7792. Girona: Museu dArqueologia de
Catalunya.
Haeussler, A. M., Turner, C. G. II & Irish, J. D.
(1988). Concordance of American and Soviet
methods in dental anthropology. Am. J. phys.
Anthrop. 75, 218.
Hanihara, K. (1967). Racial characteristics of the
dentition. J. Dental Res. 46, 923926.
Hanihara, T. (1992). Dental and cranial anities
among populations of East Asia and the Pacic: the
basic populations in East Asia, IV. Am. J. phys.
Anthrop. 88, 163182.
Haverkort, C. & Lubell, D. (1999). Cutmarks on
Capsian human remains: implications for Maghreb
Holocene social organization and palaeoeconomy.
Int. J. Osteoarch. 9, 147169.
Henke, W. (1990). Die morphometrischen anitaten
der Epipalaolithiker Nordafrikas zu populationen
Europas und des Vorderen Orients. Homo 41, 146
201.
Hershkovitz, I., Speirs, M. S., Frayer, D., Nadel, D.,
Wish-Baratz, S. & Arensburg, B. (1995). Ohalo II
H2: A 19,000 year old skeleton from a water-logged
site at the sea of Galilee, Israel. Am. J. phys. Anthrop.
96, 215234.
Hiernaux, J. (1975). The People of Africa. New York:
Charles Scribners Sons.
Holliday, T. W. (1995). Body size and proportions in
the Late Pleistocene Western Old World and the
origins of modern humans. Ph.D. Dissertation,
University of New Mexico, Albuquerque, NM.
Ighilahriz, F. (1996). LOcre dans lindustrie lithique
de lIberomaurusian dAfalou-bou-Rhummel
(Algerie). LAnthropologie 100, 7787.
Irish, J. D. (1993). Biological anities of Late Pleis-
tocene through modern African aboriginal popu-
lations: the dental evidence. Ph.D. Dissertation,
Arizona State University, Tempe, AZ.
Irish, J. D. (1997). Characteristic high- and low-
frequency dental traits in sub-Saharan African popu-
lations. Am. J. phys. Anthrop. 102, 455467.
408 . b. inisn
Irish, J. D. (1998a). Anites des populations dAfrique
du Nord et dAfrique sub-saharienne depuis la n du
Pleistoce`ne jusqua` lActuel, dapre`s la morphologie
dentaire. Bull. Mem. Soc. Anthropol. Paris 10, 237
272.
Irish, J. D. (1998b). Diachronic and synchronic dental
trait anities of Late and post-Pleistocene peoples
from North Africa. Homo 49, 138155.
Irish, J. D. (1998c). Ancestral dental traits in recent
Sub-Saharan Africans and the origins of modern
humans. J. hum. Evol. 34, 8198.
Irish, J. D. (1998d). Dental morphological indi-
cations of population discontinuity and Egyptian
gene ow in post-Paleolithic Nubia. In (J. R. Lukacs,
Ed.) Human Dental Development, Morphology, &
Pathology: A Tribute to Albert A. Dahlberg. University
of Oregon Anthropological Papers 54, pp. 155172.
Eugene: University of Oregon Press.
Irish, J. D. (1999). The Iberomaurusians: ancestors to
subsequent North African peoples or a genetic dead-
end? Am. J. phys. Anthrop. Supplement 28, 159.
Irish, J. D. & Turner, C. G. II. (1990). West African
dental anity of Late Pleistocene Nubians. II.
Peopling of the EurafricanSouth Asian triangle.
Homo 41, 4253.
Irish, J. D. & Turner, C. G. II. (1992). Further
evidence of dental discontinuity between Mesolithic
and recent Nubians. Am. J. phys. Anthrop. Supple-
ment 14, 9394.
Ishida, H. & Dodo, Y. (1997). Cranial variation
in prehistoric human skeletal remains from the
Marianas. Am. J. phys. Anthrop. 104, 399410.
Johnson, A. L. & Lovell, N. C. (1994). Biological
dierentiation at Predynastic Naqada, Egypt: An
analysis of dental morphological traits. Am. J. phys.
Anthrop. 93, 427433.
Julien, C. (1970). History of North Africa. London:
Routledge & Kegan Paul.
Konigsberg, L. W. (1990). Analysis of prehistoric bio-
logical variation under a model of isolation by geo-
graphic and temporal distance. Hum. Biol. 62,
4970.
Kruskal, J. B. & Wish, M. (1978). Multidimensional
Scaling. Beverly Hills, CA: Sage Publications.
Lahr, M. M. & Arensburg, B. (1995). Skeletal robus-
ticity in the Epipaleolithic of North Africa and the
Levant. Paleorient 21, 8796.
Lipschultz, J. G. (1996). Who were the Natuans? A
dental assessment of their population anities. M.A.
Thesis, Arizona State University, Tempe, AZ.
Lubell, D. (1984). Paleoenvironments and Epi-
Paleolithic economies in the Maghreb (ca. 20,000
5,000 B.P.). In (J. D. Clark & S. A. Brandt, Eds)
From Hunters to Farmers: The Causes and Consequences
of Food Production in Africa, pp. 4156. Berkeley:
University of California Press.
Lubell, D. (2000). Late Pleistocene-Early Holocene
Maghreb. In (P. N. Peregrine & M. Ember, Eds) The
Encyclopedia of Prehistory, Volume 1: Africa. New
York: Plenum.
Lubell, D., Sheppard, P. & Jackes, M. (1984). Conti-
nuity in the Epipaleolithic of northern Africa with
emphasis on the Maghreb. In (F. Wendorf & A. E.
Close, Eds) Advances in World Archaeology, vol. 3,
pp. 143191. New York: Academic Press.
Lubell, D., Sheppard, P. & Gilman, A. (1992). The
Maghreb. In (R. W. Ehrich, Ed.) Chronologies in Old
World Archaeology, 3rd Edn, pp. 305308 (vol. I),
257276 (vol. II). Chicago: University of Chicago
Press.
Lukacs, J. R., Hemphill, B. E. & Walimbe, S. R.
(1998). Are Mahars autochthonous inhabitants of
Maharashtra?: A study of dental morphology
and population history in South Asia. In (J. R.
Lukacs, Ed.) Human Dental Development,
Morphology, & Pathology: A Tribute to Albert A.
Dahlberg. University of Oregon Anthropological
Papers 54, pp. 119153. Eugene, OR: University of
Oregon Press.
Medig, M., Meier, R., Sahnouni, M. & Derradji, A.
(1996). Decouverte dun crane humain dans les
niveaux iberomaurusiens de la grotte de Taza I, Jijel,
Algerie. C.R. Acad. Sci. Paris 323, 825831.
Mercer, J. (1980). The Canary Islanders: Their Prehis-
tory, Conquest and Survival. London: Rex Collings.
Mourant, A. E. (1983). Blood Relations: Blood Groups
and Anthropology. Oxford: Oxford University Press.
Nurse, G. T., Weiner, J. S. & Jenkins, T. (1985). The
Peoples of Southern Africa and Their Anities. Oxford:
Clarendon Press.
Petit-Marie, N. & Dutour, O. (1987). Holocene
populations of the western and southern Sahara:
Mechtoids and paleoclimates. In (A. E. Close, Ed.)
Prehistory of Arid North Africa: Essays in Honor
of Fred Wendorf, pp. 259286. Dallas: Southern
Methodist University Press.
Philipson, D. W. (1994). African Archaeology, 2nd edn.
Cambridge: Cambridge University Press.
Pond, A. W. (1928). A Contribution to the Study of
Prehistoric Man in Algeria, North Africa. Beloit, WI:
The Logan Museum.
Roler, K. L. (1992). Near Eastern dental variation past
and present. M.A. Thesis, Arizona State University,
Temple, AZ.
Roychoudhury, A. K. & Nei, M. (1988). Human Poly-
morphic Genes World Distribution. New York: Oxford
University Press.
Sanchez-Mazas, A., Excoer, L. & Langaney, A.
(1986). Measurement and representation of the gen-
etic similarity between populations by the percentage
of isoative genes. Theoria 4, 143154.
Scott, G. R. (1973). Dental morphology: a genetic
study of American White Families and variation in
living southwest Indians. Ph.D. Dissertation, Arizona
State University, Tempe, AZ.
Scott, G. R. (1980). Population variation of Carabellis
trait. Hum. Biol. 52, 6378.
Scott, G. R. & Turner, C. G. II. (1997). The Anthro-
pology of Modern Human Teeth: Dental Morphology
and its Variation in Recent Human Populations.
Cambridge: Cambridge University Press.
Scott, G. R., Yap Potter, R. H., Noss, J. F.,
Dahlberg, A. A. & Dahlberg, T. (1983). The dental
409 incno:acncsiaN arriNi1ics
morphology of Pima Indians. Am. J. phys. Anthrop.
61, 1331.
Schwidetsky, I. (1963). La Poblacion Prehispanica de las
Islas Canarias. Santa Cruz de Tenerife: Museo
Arqueologico de Tenerife.
Sheppard, P. J. (1987). The Capsian of North Africa:
Stylistic Variation in Stone Tool Assemblages. Oxford:
B.A.R. International Series 353.
Sheppard, P. J. & Lubell, D. (1990). Early Holocene
Maghreb prehistory: An evolutionary approach.
Sahara 3, 6369.
Sjvold, T. (1973). Occurrence of minor non-metrial
variants in the skeleton and their quantitative treat-
ment for population comparisons. Homo 24, 204
233.
Sjvold, T. (1977). Non-metrical divergence between
skeletal populations. Ossa 4, 1133.
Smith, P. (1979). Regional diversity in Epipaleolithic
populations. Ossa 6, 243250.
Smith, P. & Shegev, M. (1988). The dentition of
Nubians from Wadi Halfa, Sudan: an evolutionary
perspective. J. Dent. Assoc. S. Afr. 43, 539541.
Thoma, A. (1984). Morphology and anities of the
Nazlet Khater man. J. hum. Evol. 13, 287296.
Trigger, B. G. (1976). Nubia Under the Pharaohs.
Boulder, CO: Westview Press.
Turner, C. G. II. (1967). Dental genetics and
microevolution in prehistoric and living Koniag
Eskimo. J. Dent. Res. Supplement to No. 5 46, 911
917.
Turner, C. G. II. (1985a). Expression count: a method
for calculating morphological dental trait frequencies
by using adjustable weighting coecients. Am. J.
phys. Anthrop. 68, 263268.
Turner, C. G. II. (1985b). The dental search for Native
American origins. In (R. Kirk & E. Szathmary,
Eds) Out of Asia: Peopling the Americas and the
Pacic, pp. 3178. Canberra: The Journal of Pacic
History.
Turner, C. G. II. (1987). Late Pleistocene and
Holocene population history of East Asia based
on dental variation. Am. J. phys. Anthrop. 73, 305
322.
Turner, C. G. II. (1992a). The dental bridge between
Australia and Asia: Following Macintosh into the
East Asian hearth of humanity. Persp. Hum. Biol.
2/Archaeol. Oceania 27, 120127.
Turner, C. G. II. (1992b). Microevolution of East
Asian and European populations: a dental perspec-
tive. In (T. Akaszawa, K. Aoki & T. Kimura, Eds)
The Evolution and Dispersal of Modern Humans in Asia,
pp. 415438. Tokyo: Hokusen-Sha.
Turner, C. G. II & Scott, G. R. (1977). Dentition of
Easter Islanders. In (A. A. Dahlberg & T. M. Graber,
Eds) Orofacial Growth and Development, pp. 229249.
The Hague: Mouton.
Turner, C. G. II & Markowitz, M. (1990). Dental
discontinuity between late Pleistocene and recent
Nubians. I. Peopling of the Eurafrican-South Asian
Triangle. Homo 41, 4253.
Turner, C. G. II, Nichol, C. R. & Scott, G. R. (1991).
Scoring procedures for key morphological traits of
the permanent dentition: the Arizona State Univer-
sity dental anthropology system. In (M. A. Kelley &
C. S. Larsen, Eds) Advances in Dental Anthropology,
pp. 1332. New York: Wiley-Liss.
Vallois, H. V. (1969). Les hommes de Cro-Magnon et
les Guanches: les faits acquis et les hypotheses.
Simposio del Cro-Magnon, Anuario Estudias Atlanticos,
Madrid 15, 97119.
Vaufrey, R. (1933). Notes sur le Capsien.
LAnthropologie 43, 457483.
Wendorf, F. (1968). A Nubian nal Paleolithic grave-
yard near Jebel Sahaba, Sudan. In (F. Wendorf, Ed.)
The Prehistory of Nubia, vol. 2, pp. 954995. Dallas,
TX: Fort Burgwin Research Center.
Wendorf, F. & Schild, R. (1986). Conclusion. In (A. E.
Close, Ed.) The Prehistory of Wadi Kubbaniya, vol. 1,
The Wadi Kubbaniya Skeleton: A Late Paleolithic
Burial from Southern Egypt, pp. 7174. Dallas: South-
ern Methodist University Press.
Wulsin, F. R. (1941). The Prehistoric Archaeology of
Northwest Africa. Cambridge, MA: Peabody Museum
of American Archaeology & Ethnology, Harvard
University.
Wysner, G. M. (1945). The Kabyle People. New York:
privately printed.
Zabkar, L. V. & Zabkar, J. (1982). Semna South. A
preliminary report of the 196668 excavation of the
University of Chicago Oriental Institute expedition
to Sudanese Nubia. J. Am. Res. Center Egypt 19,
2128.
410 . b. inisn