AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 000:000000 (2011)

Sahara: Barrier or Corridor? Nonmetric Cranial Traits and Biological Afnities of North African Late Holocene Populations
Efthymia Nikita,1* David Mattingly,2 and Marta Mirazon Lahr1
1

Leverhulme Centre for Human Evolutionary Studies, Department of Biological Anthropology, University of Cambridge, Cambridge CB2 1QH, UK 2 School of Archaeology and Ancient History, University of Leicester, Leicester LE1 7RH, UK KEY WORDS Garamantes; North Africa; nonmetric traits; biodistance analysis; mean measure of divergence; Mahalanobis distance ABSTRACT The Garamantes ourished in southwestern Libya, in the core of the Sahara Desert $3,000 years ago and largely controlled trans-Saharan trade. Their biological afnities to other North African populations, including the Egyptian, Algerian, Tunisian and Sudanese, roughly contemporary to them, are examined by means of cranial nonmetric traits using the Mean Measure of Divergence and Mahalanobis D2 distance. The aim is to shed light on the extent to which the Sahara Desert inhibited extensive population movements and gene ow. Our results show that the Garamantes possess distant afnities to their neighbors. This relationship may be due to the Central Sahara forming a barrier among groups, despite the archaeological evidence for extended networks of contact. The role of the Sahara as a barrier is further corroborated by the signicant correlation between the Mahalanobis D2 distance and geographic distance between the Garamantes and the other populations under study. In contrast, no clear pattern was observed when all North African populations were examined, indicating that there was no uniform gene ow in the region. Am J Phys Anthropol 000:000000, 2011. V 2011 Wiley Periodicals, Inc.
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The climate in the Sahara Desert has been hyperarid since the mid-Holocene (Cremaschi and Di Lernia, 1999; Brooks et al., 2003). Nevertheless, archaeological evidence suggests that various North African populations were in contact, exchanging material goods and cultural practices, irrespective of the harsh environment. Among others, rock art depictions of the North African Pastoral societies of the second half of the Holocene present these as multiethnic groups, suggesting some degree of intermingling with other populations (Ottoni, 2007), which is further supported by the highly mobile subsistence activities of these groups (Cremaschi and Di Lernia, 1999; Barich, 2002; Di Lernia, 2006). Naturally, the Nile River was an important corridor for population movement in North Africa. This corridor shaped relations primarily between Egypt and Nubia, whether friendly in the form of trade networks and exchange of cultural practices (Nordstrom, 1972; Gatto, 1997; Williams, 1997; Edwards, 2007) or hostile, characterized by raids on both sides (Fattovich, 1997; Davies, 2003; Edwards, 2007). Population movements across the Sahara were further facilitated by the introduction of the horse into North Africa (Mitchell, 2005). According to the distribution of rock depictions of chariots pulled by horses found in the Sahara Desert, it appears that Late Holocene trade networks emphasized the north and the northeast, with the Garamantes, a population occupying southwestern Libya, largely monitoring the trafc (Mauny, 1978; Mitchell, 2005). The Garamantes appeared in Fazzan, southwestern Libya, at $900 BC, and ourished in this area until 500 AD. What is extraordinary about them is that, although they occupied a hyperarid environment, they brought important socio-economic changes to the Central Sahara,
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such as the development of urbanism, irrigated agriculture, trans-Saharan trade, and a hierarchical, probably slave-using, society. These innovations are thought to have led to a substantial increase in population (Mattingly, 2006). Among their economic activities, trans-Saharan trade was of particular importance. Indeed, the Garamantes were involved in the trade routes that connected Fazzan with the Mediterranean littoral, Egypt, the Maghreb, and sub-Saharan Africa (Liverani, 2000). These contacts are supported by the presence of various Roman artifacts, such as glassware, ne ceramics, amphorae, cast bronze statues, architectural motifs, and masonry styles of Mediterranean tradition, in Garamantian contexts (Daniels, 1989; Mattingly, 2003a). On the other hand, the Garamantes most likely exported dates, salt, gold, semiprecious stones, jewels, and natron (Mattingly, 2003b). The Garamantes had also established contacts with other Saharan populations, as indicated by the similarities in
Grant sponsors: The Society for Libyan Studies, The Leverhulme Trust, Alexander S. Onassis Public Benet Foundation, the A.G. Leventis Foundation, and the Cambridge European Trusts (George and Marie Vergottis Cambridge Bursary). *Correspondence to: Efthymia Nikita, Leverhulme Centre for Human Evolutionary Studies, Department of Biological Anthropology, University of Cambridge, Fitzwilliam Street, Cambridge CB2 1QH, UK. E-mail: e.nikita@gmail.com Received 25 May 2011; accepted 26 October 2011 DOI 10.1002/ajpa.21645 Published online in Wiley Online Library (wileyonlinelibrary.com).

WILEY PERIODICALS, INC.

E. NIKITA ET AL.
TABLE 1. Material used in nonmetric biodistance analyses

Population-origin Garamantes Kerma Soleb Gizeh Alexandria Carthage Ain Beida, Skikda, Setif, Bijaia, Khenchela

Symbol GAR KER SOL GIZ ALE CAR ALG

Males 20 45 10 85 11 12 8

Females 26 88 17 65 9 8 12

Date 900BC500 AD 20001550 BC 15751380 BC 664343 BC 323 BC330 AD 751BC435 AD 1500 BC

Country Libya Sudan Sudan Egypt Egypt Tunisia Algeria

Collection Museum of Jarma (Libya) Duckworth Lab. (Cambridge) Musee de lHomme (Paris) Duckworth Lab. (Cambridge) Musee de lHomme (Paris) Musee de lHomme (Paris) Musee de lHomme (Paris)

their funerary structures (keyhole monuments, antenna tombs) (Mattingly, 2003c), while the technology of the subterranean irrigation channels that the Garamantes used (foggaras) was probably an Egyptian import, supporting a cultural link between the two polities (Wilson, 2003). Finally, according to written sources, the Garamantes were capturing and trading slaves from subSaharan Africa (Mattingly, 2000). On the basis of the above data, it becomes clear that North African populations were in contact throughout the Holocene. However, the hyper-arid environment of the Sahara Desert, coupled with its harsh physiography, characterized by sand seas, swampy depressions, and mountainous areas (Camps, 1982), must have restricted extensive population movements. Therefore, the material goods and cultural practices exchanged among populations during the Holocene were due to economic networks, which must have involved primarily a small subset of specialized traders. It is unlikely that such a physical barrier would have allowed long-distance marital exchanges and actual population migrations. Therefore, osteologically we would expect an increased biological distance between the Garamantes, who are geographically located at the core of the desert, and other North African populations. In addition, we would expect to nd rather high biodistances in all populations among which the desert intervened, and smaller distances among those who were geographically closer and did not have the desert intervening among them. Finally, although, as mentioned above, we do not expect to trace long-distance marriage networks, given the numerous genetic studies that have found much greater female migration rate in agricultural populations in comparison to males (for example, Seielstad et al., 1998; Destro-Bisol et al., 2004; Wooding et al., 2004; Wood et al., 2005), we would still expect females to exhibit relatively greater gene ow, thus, smaller biodistances in comparison to males.

MATERIALS AND METHODS Osteological samples

The populations used for the assessment of Garamantian biological afnities are given in Table 1, their geographic location is depicted in Figure 1 and the straightline distances between sites are shown in Table 2. Assessment of the sex and age of the individuals was based on standard osteological methods (Buikstra and Ubelaker, 1994). Given that hyperostotic traits, which constitute a large part of the current dataset, involve a progressive ossication, which may continue throughout life, subadults were eliminated from all comparisons (Buikstra, 1972). Because of lack of availability, sample sizes for Alexandria, Algeria, Soleb and Tunisia are small, especially if sexes are treated separately. As explained below, to avoid biases due to small sample American Journal of Physical Anthropology

sizes, only the statistically signicant results were taken into account in the conclusions of the present study. The Kerma skeletons were excavated in the Eastern Cemetery by Reisner in 19131914 and 19151916, and date between 2000 and 1550 BC (Keita, 1988; Filer, 1992; Zakrzewski, 2003). Kerma was the capital of the Kingdom of Kush in Upper Nubia (Hafsaas-Tsakos, 2009). This population controlled trade between Egypt, central Africa and the East, and was characterized by social stratication (Adams, 1977). In addition, Kermas position on a oodplain along the Upper Nubian Nile enabled productive agriculture and exploitation of the local mineral resources (Trigger, 1976; Adams, 1977). Nubian skeletons from the site of Soleb date to 1575 1380 BC (Trigger, 1976). During this period, Nubia was under Egyptian control, the state of Kerma had been destroyed and the viceroy of Kush took over the general administration moving the capital in Miam (Aniba), although the local administration was in the hands of tribal leaders (Adams, 1977). The Gizeh material used in the present study belongs to the so-called E Series, and was excavated by Petrie in a cemetery located to the south of the Gizeh pyramids. It is dated to 664343 BC (Petrie, 1907), a period characterized by intermittent Persian occupation (Lloyd, 2000). Ptolemaic and Roman skeletons from Alexandria (323 BC330 AD) were also examined. Alexandria controlled trade across the Mediterranean, as well as with India and the East African coast (Phillipson, 2005), and exploited the inland as a source of agricultural goods and natural resources (Haas, 1997). In the Roman period, its population uctuated between 500,000 and 600,000 individuals (Delia, 1988), although others suggest a lower estimate (just over 300,000) (Alston and Alston, 1997). Social differentiation was present during that period, with Greek and urban representing the highest status, while Egyptian and rural the lowest. This stratication resulted in increased ethnic tensions and a series of revolts (Alston, 1997). The Phoenician skeletons are from Carthage, Tunisia, and date to the Punic and early Roman period (751 BC435 AD) (Wysner, 1945; Charles-Picard and Picard, 1968). The Phoenician civilization ourished across the Mediterranean by establishing widely distributed commercial outposts and was organized in city-states. Aside from trading, their economy was based on cereal cultivation (Phillipson, 2005). Among the comparative North African populations were also some Algerians (1500 BC) (Keita, 1988). The origin of these groups is uncertain, but they were most likely autochthonous to the region (Camps, 1974; Lefevre-Witier et al., 2006).

METHODS
The biological afnities of the various populations were examined by means of cranial and mandibular

SAHARA: BARRIER OR CORRIDOR?

Fig. 1. Selected archaeological sites within the region of study. TABLE 2. Straight-line geographic distances in km among populations under study Garamantes Garamantes Algerians Alexandrians Gizeh Kerma Soleb Carthaginians 1,235 1,730 1,810 1,920 1,860 1,190 Algerians 2,180 2,350 2,900 2,850 260 Alexandrians Gizeh Kerma Soleb

180 1,280 1,185 1,950

1,150 1,070 2,110

90 2,770

2,690

nonmetric traits. Nonmetric traits are believed to reect an expression of genes that affect development and follow a non-Mendelian pattern of inheritance (Berry and Berry, 1967; Ossenberg, 1984; Hauser and De Stefano, 1989; Tyrrell, 2000). Twin and family studies, as well as studies on laboratory animals, have demonstrated that the majority of these traits are under strong genetic control (Saunders and Popovich, 1978; Cheverud and Buikstra, 1981a, 1981b, 1982; Carson 2006). However, environmental factors also have some inuence as trait expression seems to follow the threshold model proposed by Falconer (Gruneberg, 1952). In any case, there is sufcient genetic control to make nonmetric traits useful in anthropological work, and such variables have successfully assigned individuals from distinct geographic regions to their correct groups (for example, Winder, 1981; Prowse and Lovell, 1995; Blom et al., 1998; Christensen, 1998; Donlon, 2000; Hanihara et al., 2003). As such, nonmetric traits are an appropriate means for studying biological distances among North African populations and the implications of the patterning of these distances in assessing the relative role of the Sahara Desert as a barrier to gene ow. The 34 traits employed in the current study, along with their scoring details, are given in Table 3. Note that for cranial traits, no standard protocol exists that denes where the threshold between presence and absence should be placed and this lack may have an important impact on the obtained results (Carson, 2006). For most traits, the criteria employed by Dodo (1974) were used. The remaining relevant thresholds were dened by Nikita (2010) as presented in Table 3. The traits were selected on the basis of their successful use by other researchers performing biodistance analyses (Prowse

and Lovell, 1996; Hanihara et al., 2003; Sutter and Mertz, 2004). Each trait was scored as present/absent, and its frequency was calculated based on the individual count method, that is, ignoring potential bilateral presence and only indicating whether the individual shows the trait (Sutter and Mertz, 2004). This method was used because unilateral scoring of cranial nonmetric traits reveals a strong correlation between sides (McGrath et al., 1984). The side with the highest degree of expression was used (Hauser and De Stefano, 1989; Scott and Turner, 1997). While it is recognized that poor sample preservation has the potential to bias the results obtained (Konigsberg, 1987), the remains analyzed in the current paper were well preserved overall. After the initial scoring, sex dimorphism in trait expression was explored with chi-square analysis. The statistical package used was SPSS for Windows, version 18.0. Furthermore, nondiscriminatory traits across samples were removed. In particular, following the recommendation of Harris and Sjvold (2004), all traits that did not exhibit at least one statistically signicant difference between sample pairs were removed from the data set. Statistical analyses used for this purpose were Pearsons chi-square tests or Fishers exact tests run in SPSS. In addition, all associated traits were identied using Pearsons chi-square test or Fishers exact test (Irish, 2010 and references therein). For that purpose a homemade macro was written in Excel Visual Basic for Applications (VBA) that identied the traits which exhibited the most numerous associations, removed them from the dataset one by one and repeated the analyses until no associated traits remained. Two measures for the assessment of pairwise biodistances among the various samples were adopted after American Journal of Physical Anthropology

E. NIKITA ET AL.
TABLE 3. Nonmetric cranial traits used in this study and references

Trait name Metopic suture Metopic ssure Supranasal suture Supraorbital osseous structures Divided infraorbital foramina Parietal foramina Divided mental foramina Ethmoidal foramina Lesser palatine foramina Squamous ossicles Parietal notch bone Epipteric bone Ossicle at asterion Occipitomastoid wormians Coronal ossicles Sagittal ossicles Lambdoid ossicles Inca bone Divided occipital condyles Hypoglossal canal bridging Mandibular torus Maxillary torus Auditory torus Palatine torus Apertures at the oor of the acoustic meatus Divided parietal bone Divided temporal squama Os japonicum Marginal tubercle Mylohyoid bridging Foramen of Vesalius Foramen ovale incomplete Zygomaxillary tubercle Symmetrical thinness of parietal bones

Description Hauser and De Stefano (1989) Hauser and De Stefano (1989) Hauser and De Stefano (1989) Dodo (1974) Hauser and De Stefano (1989) Hauser and De Stefano (1989) Hauser and De Stefano (1989) Hauser and De Stefano (1989) Hauser and De Stefano (1989) Hanihara and Ishida (2001) Dodo (1974) Dodo (1974) Ossenberg (1969) Dodo (1974) Hanihara and Ishida (2001) Hanihara and Ishida (2001) Dodo (1974) Dodo (1974) Hauser and De Stefano (1989) Dodo (1974) Dodo (1974) Hauser and De Stefano (1989) Kennedy (1986) Hauser and De Stefano (1989) Hauser and De Stefano (1989) Hauser and De Stefano (1989) Hauser and De Stefano (1989) Dodo (1974) Hauser and De Stefano (1989) Dodo (1974) Dodo (1974) Dodo (1974) Hauser and De Stefano (1989) Hauser and De Stefano (1989)

Presence threshold Present along more than half of the frontal arc (Dodo 1974) Present in any of its variables (see Hauser and De Stefano 1989) If observable irrespective of degree of persistence or shape No distinction between notches and foramina, present if open to the orbital cavity (Dodo 1974) Only complete bridging If observable irrespective of position, size or number Only complete division If the posterior foramen is absent If observable irrespective of position, size, shape or number If observable irrespective of size or number (Hauser and De Stefano 1989) If observable irrespective of predominant position, size or number (Dodo 1974) If observable irrespective of size, type of articulation with neighboring bones or number If observable irrespective of predominant position, size, shape or number (Ossenberg 1969) If observable irrespective of exact position, size or number (Dodo 1974) If observable irrespective of position, size or number If observable irrespective of position, size or number If observable irrespective of position, size or number If more than 10mm of the suture remained (Dodo 1974) If a deep furrow cutting into the facet from both sides could be seen, even if it did not completely separate the condyle (Hauser and De Stefano 1989) Only complete division (Dodo 1974) Any expression from weak to strong (Dodo 1974) Any expression from weak to strong Any expression from weak to strong Any expression from weak to strong At least pinhole sized apertures (Hauser and De Stefano 1989) If observable for more than 1 cm If observable for more than 5mm If observable for more than 5mm (Dodo 1974) If projecting for more than 4 mm If osseous bridge observable irrespective of location and degree of expression (Dodo 1974) Only complete divisions Any kind of communication between the two foramina except for a suture-like gap (Dodo 1974) If projecting for more than 2 mm When ranging from slight attening to a saucer-shaped appearance (Hauser and De Stefano 1989)

trait editing: the modied Smiths Mean Measure of Divergence (MMD) (Sjvold, 1973) and the generalized Mahalanobis distance (D2) utilizing a tetrachoric correlation matrix for nonmetric data (Konigsberg, 1990). Although MMD has been criticized by several scholars on various grounds (Konigsberg and Buikstra, 2006; Harris, 2008), it still remains the most popular statistic used in anthropological studies (see for example, Green and Suchey, 1976; De Souza and Houghton, 1977; Smith, 1977; Finnegan and Cooprider, 1978; Irish, 1998; Hanihara et al., 2003; Sutter and Mertz, 2004; Ossenberg et al., 2006; Pilbrow, 2006; Irish, 2010 and discussion therein). The MMD was estimated from (Sjvold, 1973):  r  1X 1 1 MMD 1 u1i u2i 2 r i1 n1i 0:5 n2i 0:5 where r is the number of traits considered, n1i is the number of crania in population 1 examined for trait i, n2i is the number of crania in population 2 examined for American Journal of Physical Anthropology

trait i and y1i and y2i are the transformed frequencies. As such, the MMD is the average value of the Measure of Divergence (MD) calculated over all traits. The transformed frequencies can be estimated using various methods: the Smith (devised for use rst by Grewal, 1962), the Freeman-Tukey (1950) and the Anscombe (1948) transformations, as well as Bartletts correction (Bartlett, 1937; Green and Suchey, 1976). In the present study the Freeman-Tukey transformation was adopted since, according to Green and Suchey (1976), it stabilizes the variance more quickly when small samples of the size often considered by archaeologists are used. Thus, y values were calculated from:     1 k 1 k1 u sin1 1 2 sin1 1 2 2 2 n1 2 n1 where sin21 is the arcsin function and k is the times that a trait occurs in a sample of size n. However, in order to examine whether small frequencies (\10%) in

SAHARA: BARRIER OR CORRIDOR? the samples under study with small sizes bias the MMD values (Green and Suchey, 1976), the Anscombe transformation and Bartletts correction were also used as a preliminary control. In the cases where negative MMD values were obtained, these were converted to 0 (Harris and Sjvold, 2004; Irish, 2006; Sutter and Verano, 2007; Irish, 2010). At this point we should clarify that negative MMD values imply that the biological distance between two populations is very small. Indeed, when two populations are very close, then y1i % y2i and, therefore, a negative contribution to MMD may arise from the term 1=n1i 0:5 1=n2i 0:5: The variance of the MMD was calculated from (Sjvold, 1973): 2 r  2X 1 1 3 VarMMD 2 r i1 n1i 0:5 n2i 0:5 p In general, if MMD= VarMMD [ 2, the compared samples are regarded as signicantly divergent at the 0.025 level (Sjvold, 1977). Nevertheless, in the present paper the p values for each MMD were estimated using two methods. The rst method was based on the fact that the MMD values follow the normal distribution (Sjvold, 1973; Green et al., 1979) and the second one used permutations (10,000 in the current study). The generalized Mahalanobis D2 distance for nonmetric data proposed by Konigsberg (1990) has been used by several scholars (for example, Bedrick et al., 2000; Sutter and Verano 2007; Schillaci et al., 2009; Irish 2010). This distance was calculated from: D2 zik zjk 0 T1 zik zjk ij
21

TABLE 4. Signicant sex differences in nonmetric trait frequencies Pearson Fishers Chi-square exact test P value P value 0.042 0.025 0.041 0.005 0.000 0.044 0.015 0.033 0.044 0.001 0.007 0.011 0.001 0.020a

Trait Perietal foramina Ethmoidal foramina Hypoglossal canal bridging Supranasal suture Divided mental foramina Epipteric bone

Population Garamantes Garamantes Garamantes Garamantes Gizeh Carthaginians Kerma

Kerma Gizeh Apertures at the oor Kerma of the acoustic meatus Gizeh Marginal tubercle Kerma Zygomaxillary tubercle Kerma Gizeh Palatine torus Gizeh
a

0.040a

Sample sizes \ 5.

represents the inversed pooled tetrachoric where T correlation matrix between k traits, and zik and zjk are threshold values related to trait k in samples i and j, respectively. The signicance of D2 was estimated using ij the following variable (Sutter and Verano, 2007): Ngr1 ni nj D2 F r ni nj N g ij 5

between (a) the MMDs of males and females, (b) the MMD and D2 matrices for pooled sexes, and (c) the biological and geographic distance of the populations under study were examined. In addition, partial Mantel tests were run to explore the correlation between biological distance and geographic distance, controlling for the effects of temporal differences among populations. All Mantel tests were performed in R (functions mantel and mantel.partial of the vegan library). The correlations between biological distance and geographic distance were also explored using bootstrap correlation in R language (functions boot and boot.ci of the boot library). The number of permutations was set to 50,000.

RESULTS Sexual dimorphism

The application of Pearsons chi-square test or Fishers exact test (depending on the sample sizes) demonstrates that, for certain variables, signicant differences in trait frequency occur between the two sexes (Table 4). However, none of the variables differs consistently in frequency between males and females in all populations, and most exhibit signicant differences only in one out of the seven samples under examination. Nevertheless, the male/female ratio ranges between 0.4 and 1.5 in the current samples and this is an indication that sexes may be treated separately. For this reason, all interpopulation comparisons were initially performed separately for males and females. However, since the effect of the above sex differences on the biodistances is not known a priori, especially after trait editing, this issue is re-examined on the basis of correlations between biodistance matrices in the section Population afnities.

where N is the total number of individuals, g is the number of populations, r is the number of traits, and ni and nj is the number of individuals in populations i and j, respectively. This variable follows the F distribution with r and (N g r 1 1) degrees of freedom. The generalized Mahalanobis D2 distance was estimated using a home-made macro written in Microsoft Excel Visual Basic for Applications, where the tetrachoric matrix was calculated based on the algorithm AS 116 (Brown, 1977). To visualize intersample phenetic afnities, each MMD and D2 matrix was analyzed by conducting interval-level metric multidimensional scaling (MDS) using SPSS 18.0 Procedure Alscal, following Irish (2010). To assess which individual traits are driving the patterns observed in MDS, the Measure of Divergence of each trait in each pairwise comparison was calculated from: 1 1 6 n1i 0:5 n2i 0:5 Mantel tests were performed to examine differences between distance matrices. In particular, the correlations MDi u1i u2i 2

Trait editing
Pearsons chi-square test and Fishers exact test showed that 21 traits among males and another 21 traits among females did not differ signicantly in any of the pairwise comparisons among populations. Consequently, all these traits were dropped. The remaining traits included the metopic and supranasal sutures, supraorbital structures, American Journal of Physical Anthropology

E. NIKITA ET AL.
TABLE 5. Trait frequencies among populations under study GAR ALG 33.3 83.3 0.0 66.7 16.7 0.0 0.0 33.3 33.3 66.7 50.0 91.7 8.3 50.0 41.7 16.7 11.3 16.7 33.3 25.0 ALE 72.7 100.0 9.1 45.5 27.3 0.0 45.5 11.1 72.7 54.5 22.2 88.9 0.0 55.6 66.7 22.2 0.0 0.0 55.6 22.2 GIZ 85.9 97.6 2.4 89.4 21.2 10.6 61.2 37.6 60.0 71.8 55.4 100.0 3.1 60.0 86.2 36.9 10.8 24.6 52.3 44.6 KER 84.4 95.6 0.0 64.4 6.7 0.0 53.3 20.0 82.2 80.0 31.8 90.9 0.0 69.3 79.5 23.9 18.2 19.3 54.5 58.0 SOL 70.0 100.0 0.0 40.0 20.0 0.0 30.0 10.0 30.0 40.0 17.6 88.2 8.9 35.3 58.8 23.5 0.0 11.8 11.8 35.3 CAR 75.0 100.0 0.0 83.3 9.3 9.3 25.0 25.0 33.3 41.7 12.5 100.0 12.5 75.0 37.5 12.5 0.0 0.0 50.0 50.0

Males Supranasal suture Supraorbital structures Infraorbital foramina Ethmoidal foramina Epipteric bone Coronal ossicles Lambdoid ossicles Hypoglossal canal bridging Marginal tubercle Zygomaxillary tubercle Females Supranasal suture Supraorbital structures Infraorbital foramina Parietal foramina Ethmoidal foramina Epipteric bone Palatine torus Apertures at the oor of auditory meatus Marginal tubercle Zygomaxillary tubercle

65.0 85.0 30.0 35.0 0.0 0.0 25.0 0.0 30.0 30.0 23.1 69.2 11.5 38.5 26.9 3.8 0.0 3.8 38.5 7.7

infraorbital and ethmoidal foramina, epipteric bone, coronal and lambdoid ossicles, hypoglossal canal bridging, mandibular torus, marginal tubercle, fused ovale and spinous foramina, and zygomaxillary tubercle for males. For females, the remaining traits were ossicle at asterion, epipteric bone, mandibular and palatine tori, fused ovale and spinous foramina, infraorbital, parietal and ethmoidal foramina, apertures at the oor of the auditory meatus, marginal and zygomaxillary tubercle, supranasal suture, and supraorbital structures. Assessment of associated traits led to further removal of the metopic suture, the mandibular torus, and the fused ovale and spinous foramina in males, and the ossicle at asterion, the mandibular torus, and the fused ovale and spinous foramina in females. The frequencies of the remaining traits are given in Table 5. At this point it is interesting to note that when the above edited data are used for the calculation of MMD, the P values estimated using permutations agree with those calculated using the normal distribution (Fig. 2A). In contrast, when the unedited data (all 34 traits) are analyzed, the P values obtained by the two methods are different (Fig. 2B). These results suggest that when the editing of the traits is correct and associated as well as nondiscriminatory variables are eliminated, the MMDs indeed follow a normal distribution. Therefore, the overlap of the P values estimated by the two methods described above may be used as a criterion to check whether data editing is successful or not.

Population afnities
Table 6 presents the MMD values for males and females estimated using Freeman-Tukey transformation. As already pointed out, some of the samples under study exhibit small sizes, particularly when males and females are studied separately. In principle, statistics does not preclude the examination of small samples; however, in any statistical test involving small sample sizes, only the rejection of the null hypothesis is of value. For this reason, the conclusions of this study will only take into consideration the statistically signicant results, i.e., when the null hypothesis is rejected. Furthermore, it should be American Journal of Physical Anthropology

Fig. 2. Variation of P-value in inter-population comparisons for males using the edited (A) and the original data (B). Values have been estimated using permutations (~) and the normal distribution (D).

stressed that an additional issue when small sample sizes are analyzed using nonmetric traits is the possible estimation of wrong MMD values. As Green and Suchey (1976) have shown, such wrong estimates occur when the frequency of traits is small (\10%). However, as can be seen in Table 5, in this study very few traits exhibited such small frequencies in the small sized populations. Therefore, the calculated MMDs are likely to be unbiased

SAHARA: BARRIER OR CORRIDOR?

TABLE 6. Biological distances using MMD for males and females GAR GAR ALG ALE GIZ KER SOL CAR 0 0.211* 0.108 0.435*** 0.345*** 0.063 0.165* ALG 0.079 0 0.117 0.212** 0.221** 0.029 0 ALE 0.005 0 0 0.113* 0.036 0 0.070 GIZ 0.472*** 0.001 0.084 0 0.090*** 0.170** 0.041 KER 0.400*** 0.027 0.037 0.056*** 0 0.145* 0.156** SOL 0.085* 0 0 0.209*** 0.187*** 0 0

CAR 0.072 0 0 0.122* 0.067 0.002 0

Male MMD values are given below the diagonal and female values above the diagonal. * MMD values are signicantly different at P \ 0.05, **signicant at P \ 0.01, ***signicant at P \ 0.001.

Fig. 4. Three-dimensional MDS based on the MMD values for male data. Fig. 3. Variation of MMD distance in inter-population comparisons for males (top) and females (bottom). The MMD values have been calculated using the Freeman-Tukey transformation (l), the Anscombe transformation (3) and Bartletts correction (h). Data for males have been shifted along the y-axis by 0.5 to make the picture clearer.

by small sample sizes. This estimation is further conrmed by comparing MMD values obtained by FreemanTukey transformation to those calculated from the Anscombe transformation and Bartletts correction. Figure 3 shows that the transformation method does not practically affect the MMD values. Table 6 shows that the greatest distance of the Garamantes to their neighbors is that with Gizeh and Kerma, both for males and females. These distances are also the greatest among all populations under study and they are statistically signicant. The smallest biodistance for male Garamantes is with the population from Soleb, whereas for females it is with the Alexandrians. In what concerns the general pattern of the observed distances in all populations, it is similar for males and females but females overall exhibit smaller pairwise MMDs than males (mean male MMD 5 0.13, mean female MMD 5 0.09), although this result may be biased by the small sample sizes for some groups. In any case, the above difference in mean values between sexes is not statistically signicant (Mann-Whitney test, P 5 0.096).

Multi-dimensional scaling (MDS) offers a good visualization of intersample phenetic afnities. When it is conducted using three dimensions, the obtained stress value for males is 0.058 (Kruskals stress formula 1) and r2 5 0.972, which suggests a good representation of the MMD distances. For females these results deteriorate to some extent since the stress value is equal to 0.110 and r2 5 0.884. The three-dimensional plots of the MDS analyses are shown in Figures 4 and 5. To gain a better understanding of sample biodistances, these gures have been drawn using the same scale for all axes since the range of the MDS values from Dimension 1 to 3 follows roughly the ratio 4.6: 2.5: 1.6. Figures 4 and 5 show that among males and females the Garamantes are located at the very positive extreme of Dimension 1, Gizeh and Kerma are at the opposite extreme, while all other groups are spread close to the center. With regard to Dimensions 2 and 3, the Garamantes do not seem to differ particularly from the other populations, being located close to the middle of the comparative range of values. Overall, however, the Garamantes stand out in multidimensional space, as do the samples from Kerma and Gizeh. To examine the relative position of males and females, we plotted the rst two dimensions of the MDS using again the same scale for the two axes (Fig. 6A). In this gure, we observe even more clearly the separation of the Garamantes on one hand and the Kerma and Gizeh American Journal of Physical Anthropology

E. NIKITA ET AL.

Fig. 5. Three-dimensional MDS based on the MMD values for female data.

Fig. 7. Plot of MD vs. all edited traits concerning the pairwise comparisons between male (A) and female (B) Garamantes-Gizeh (1), Garamantes-Kerma (3), Garamantes-Soleb (o) and Gizeh-Kerma (l). Traits: 1. supranasal suture, 2. supraorbital structures, 3. infraorbital foramina, 4. ethmoidal foramina, 5. epipteric bone, 6. marginal tubercle, 7. zygomaxillary tubercle, 8. coronal ossicles (males)/parietal foramina (females), 9. lambdoid ossicles (males)/ palatine torus (females), 10. hypoglossal canal bridging (males)/ apertures at the oor of auditory meatus (females).

Fig. 6. Plot of Dimension 1 vs. Dimension 2 of the threedimensional MDS (A) for males (l) and females (o), and (B) for pooled sexes.

on the other in comparison to the remaining groups along Dimension 1. Furthermore, we observe that in the samples from Gizeh, Kerma, and Garamantes males and females are close to each other. In contrast, males and females do not cluster very closely in the remaining populations. To examine which traits drive the biodistances among populations, we calculated the measure of divergence (MD) for each trait (Fig. 7). This gure depicts the MD values for all edited traits concerning the pairwise comparisons between male (A) and female (B) GaramantesAmerican Journal of Physical Anthropology

Gizeh, Garamantes-Kerma, Garamantes-Soleb, and Gizeh-Kerma. The Garamantes-Gizeh and GaramantesKerma exhibit the greatest distances along Dimension 1 in the MDS plots, whereas the Soleb sample shows the smallest distance from the Garamantes along this dimension. The MDs of Gizeh-Kerma are included because these two populations are located at the opposite extreme to the Garamantes along Dimension 1. We observe that for males the traits that drive the biodistances are mainly the infraorbital foramina and secondarily the marginal tubercle and the zygomaxillary tubercle, while for females these traits include the ethmoidal foramina, and secondarily the supraorbital structures and zygomaxillary tubercle. An interesting nding is that between the Kerma and Gizeh all traits show similar and small MDs, despite their relatively long distance in Dimension 2 (Fig. 6). It is seen that so far only small overall differences have been observed between males and females. For this reason Mantel tests were performed between the MMD distance matrices of males and females to examine whether these matrices are statistically signicantly different and subsequently, whether it would be valid to pool the sexes, despite the differences already found in the frequency of certain nonmetric traits. The Mantel tests showed a signicant correlation (Pearsons r 5 0.788, P 5 0.005). It is interesting to note that if the comparison is restricted to statistically signicant MMD

SAHARA: BARRIER OR CORRIDOR?

TABLE 7. Biological distances among the populations under study for pooled sexes GAR GAR ALG ALE GIZ KER SOL CAR 0 0.095* 0.145** 0.538*** 0.410*** 0.131*** 0.187** ALG 2.693** 0 0 0.088** 0.042 0 0 ALE 5.073*** 2.689** 0 0.108** 0.048* 0.054 0 GIZ 7.804*** 2.348*** 3.296*** 0 0.068*** 0.265*** 0.051* KER 6.903*** 3.947*** 4.022*** 5.154*** 0 0.197*** 0.050* SOL 5.545*** 3.026** 2.110** 4.525*** 9.102*** 0 0.021 CAR

2.578*** 1.582 4.424*** 7.633*** 5.153*** 3.698*** 0

MMD distances are given below the diagonal and Mahalanobis distances above the diagonal. * Distances are signicantly different at P \ 0.05, **signicant at P \ 0.01, ***signicant at P \ 0.001.

Fig. 8. Three-dimensional MDS based on the MMD values for pooled data.

Fig. 9. Three-dimensional MDS based on the Mahalanobis distances D2 for pooled data.

values, then Pearsons r increases to 0.984, with P 5 0.0004. These ndings show that males and females can be pooled for the estimation of biodistances. Table 7 gives the MMD values obtained after pooling males and females using the common nonmetric traits between sexes. MDS was also applied to this data and the three-dimensional representation of the results is given in Figure 8 (Kruskals stress 5 0.096, r2 5 0.934). The plot of Dimensions 1 and 2 of the three-dimensional MDS plot is given in Figure 6B. We observe that both plots are strikingly similar to the corresponding plots when the two sexes were studied separately. Table 7 also presents the Mahalanobis D2 distances for pooled sexes. Note that all D2 values are statistically signicant with P most often \0.001, with the only exception the distance between the Algerians and Carthaginians (P 5 0.14). The D2 values were also visualized by means of MDS and the three-dimensional plot is given in Figure 9 (Kruskals stress 5 0.068, r2 5 0.935). It is seen that again the Garamantes lie on one extreme and the Gizeh and Kerma on the opposite extreme of Dimension 1. The only difference to the MMD data is that now the Carthaginians appear closer to the Garamantes. The convergence between the two biodistance measures, MMD and D2, was statistically veried by Mantel tests. The correlation between MMD and D2 distances turned out to be signicant (Pearsons r 5 0.567, P 5 0.015). Finally, in order to examine the isolation by geographical distance, Mantel tests (Zakrzewski, 2007; Irish, 2010)

as well as bootstrap correlations (Schillaci et al., 2009) were adopted. All correlations between biological distance and linear geographic distance failed to reach statistical signicance, whether using MMD or Mahalanobis D2 (MMD vs. geographic distance: Pearsons r 5 20.162, P 5 0.843, D2 vs. geographic distance: Pearsons r 5 20.027, P 5 0.545). These values are not affected by the MMD values that did not exhibit statistical signicance. Indeed, if the nonsignicant MMD values are removed from the dataset, we obtain Pearsons r 5 20.044, P 5 0.881. The bootstrap analysis gave symmetric curves for the distribution of the correlation coefcient and condence intervals that conrmed the above results. In contrast, the examination of the correlation of biological distance to geographic distance between the Garamantes and their neighbors showed a signicant correlation for the Mahalanobis D2 distance (Pearsons r 5 0.911, P 5 0.004). For the MMD the correlation is positive but not statistically signicant (Pearsons r 5 0.511, P 5 0.165). This correlation is visualized in Figure 10, which shows that the positive correlation between MMD and geographic distance is in fact screened by the scatter of the MMD points. It should be mentioned that River and Mediterranean coast distances gave similar results to linear distances (results not shown). Concerning temporal effects, the partial Mantel tests showed that the temporal differences among populations do not affect the correlation between biological and geographic distance (MMD vs. geographic distance: PearAmerican Journal of Physical Anthropology

10

E. NIKITA ET AL. fact that the distances among the populations under study are not equivalent to each other. Thus, traveling across the Sahara desert is clearly much more difcult than traveling along the Nile River or the Mediterranean coast. Another factor that could affect the present results, as well as the results of similar studies, is the fact that isolated populations could have experienced substantial gene ow with populations not included in the analysis. However, the impact of this factor cannot be assessed. The fact that not all populations under study were strictly contemporaneous is unlikely to affect the results since the temporal effect is included in the treatment. Note that in the literature there are rather contradictory and puzzling results. For example, if we consider only populations across the Nile River, Zakrzewskis (2007) study also failed to accept an isolationby-distance model using the Mahalanobis distance for metric data. The same was veried by Schillaci et al. (2009) but, in contrast to D2, they found that the MMD distances are correlated with both temporal and straight-line geographic distances. In this study the lack of correlation between biodistance and geographic distance when all populations are considered relates to a number of unexpected results. More specically, Algerians and Carthaginians, which were geographically close to each other, also exhibited a close biological relationship (MMD 5 0, D2 5 1.58). However, exceptions to this rule have been identied, as for example the Gizeh and Alexandrians (MMD 5 0.11, D2 5 3.3) or the Kerma and Soleb (MMD 5 0.2, D2 5 9.1). Problems also appear for geographically distant populations. For example, the low biodistances between the Alexandrians and Soleb (MMD 5 0.05, D2 5 2.1) may be attributed to the long relationships between Egypt and Nubia and the fact that the Nile River formed a corridor that facilitated population contacts (Williams, 1997; Irish, 2006; Edwards, 2007; Schillaci et al., 2009). It is, however, striking and difcult to explain why this link apparently excluded the Gizeh and Soleb (MMD 5 0.27, D2 5 4.5). Our study suggests that when the Sahara desert intervenes among populations, their biodistances, MMD and D2, as a rule increase. In contrast, no clear pattern is observed for populations that could follow routes along the Nile River or/and the Mediterranean coast. Thus, we see that the Sahara desert inuenced gene ow among North African populations and inhibited extended contacts in the Late Holocene. The trade networks evidenced by the archaeological data involved a subset of each population, whereas the presence of local traders, who formed a chain among widely dispersed groups, cannot be excluded. In any case, substantial gene ow among the involved populations was obstructed. Temporal effects in distance correlations were not detected. This result is to be expected because the populations under study were chosen so that they exhibited the smallest possible temporal distances in order for the temporal effect on the biodistance results to be minimal. Another interesting point to be noted is that the relations among men and women are slightly different and might throw light on gender biases in marriage patterns. More specically, a signicant correlation was found between the biological distance matrices of males and females. This correlation suggests that both sexes engaged similarly in encounters with their neighbors. The Garamantes represent a typical example and it is possible that Garamantian females, due to their location

Fig. 10. Plot of MMD (o) and D2 (x) vs. geographic distance between the Garamantes and their neighbors. Regression equations: y 5 0.0003x 0.2074, R2 5 0.261, P 5 0.165 for MMD and y 5 0.006x 2 4.6354, R2 5 0.829, P 5 0.004 for D2.

sons r 5 20.168, P 5 0.852, D2 vs. geographic distance: Pearsons r 5 20.034, P 5 0.564).

DISCUSSION
This study aimed at assessing the extent to which the Sahara Desert was a barrier to gene ow over hundreds of years among Late Holocene North African populations. The results obtained from multidimensional scaling show an interesting pattern of afnities and differences in the region under study. Overall, three clusters may be identied. These consist of (1) the Garamantes, (2) a group formed by the populations of Gizeh and Kerma, and (3) a group that includes Soleb, Alexandrians, Algerians, and Carthaginians. The above results have important implications for our understanding of population history and afnities in North Africa in the last 4,000 years. Overall, our ndings indicate that the Garamantes, the population at the geographic center of the various trade networks, who also occupied the harshest environment with the desert intervening between them and all other groups under study, were rather isolated. This relative isolation may be the result of the increased desertication of the Central Sahara. It should be noted that both sexes appeared equally isolated. What is of particular interest in terms of the role of the Sahara Desert as a barrier in the Late Holocene is the fact that the biological distance of the Garamantes to their neighbors exhibited a signicant correlation to the corresponding geographic distance when using Mahalanobis D2. The MMD of the population from its neighbors also increased with increasing geographic distance but this correlation was not statistically signicant due to the large scatter of the MMD values. Note that Konigsberg (1990) was the rst to formally develop models of genetic isolation by geographic and temporal distance and has shown that if there is isolation by geographic distance, then the correlation between genetic and spatial distance should be positive. Therefore, our results suggest an isolation-bydistance model when the Garamantes are examined in relation to their neighbors. In contrast, when all North African populations were examined, the correlations between biodistance and geographic distance failed to accept an isolation-by-distance model irrespective of whether MMD or D2 data were considered. This result implies that there is no uniform gene ow throughout North Africa, possibly due to the American Journal of Physical Anthropology

SAHARA: BARRIER OR CORRIDOR? at the desert core, were prevented from being actively involved in long-range marital networks. On the other hand, females generally exhibited smaller pairwise biodistances compared with males, several of which were zero. However, these results are tentative since many of the biodistances (Table 6) are not statistically signicant to conrm this interesting observation, which possibly results from the greater mobility of females as part of the various marital networks. Note that this greater mobility is also in accordance to genetic data obtained from other agricultural populations (Seielstad et al., 1998; Destro-Bisol et al., 2004; Wooding et al., 2004; Wood et al., 2005), where patrilocality is the general pattern. To sum up, this study identied three broad clusters among North African Late Holocene populations. The rst consisted of the Garamantes, the second included the Gizeh and Kerma, and the third comprised of Soleb, Alexandrians, Algerians, and Carthaginians. The Garamantes, although actively involved in various transSaharan trade networks according to archaeological data, appear rather isolated and their Mahalanobis D2 biodistance to the other North African groups correlated signicantly to the corresponding geographic distances. This result most likely relates to the Sahara desert which intervened between them and their neighbors. In contrast, the study of all North African populations failed to accept an isolation-by-distance model, which suggests that there was no uniform gene ow across North Africa. In general, whenever the Sahara desert intervened among populations, their biodistances increased, whereas no clear pattern was observed for populations that could follow routes along the Nile River or/and the Mediterranean coast. Therefore, the Sahara desert restricted gene ow among North African populations and the trade networks evidenced by archaeological studies possibly involved a subset of male merchants and local traders.

11

ACKNOWLEDGMENTS
Special thanks should be addressed to all archaeologists and experts who participated in the Desert Migrations Project.

LITERATURE CITED
Adams WY. 1977. Nubia. Corridor to Africa. Princeton, NJ: Princeton University Press. Alston R. 1997. Philos In Flaccum: ethnicity and social space in Roman Alexandria. Greece Rome 44:165175. Alston R, Alston RD. 1997. Urbanism and the urban community in Roman Egypt. J Egypt Archaeol 83:199216. Anscombe FJ. 1948. The transformation of Poisson, binomial, and negative-binomial data. Biometrika 35:246254. Barich B. 2002. Cultural responses to climatic changes in North Africa: beginning and spread of pastoralism in the Sahara. In: Hassan F, editor. Droughts, food and culture. Ecological change and food security in Africas later prehistory. New York: Kluwer Academic/Plenum Publishers. p 209223. Bartlett MS. 1937. Properties of sufciency and statistical tests. Proc R Soc A 160:268282. Bedrick EJ, Lapidus J, Powell JF. 2000. Estimating the Mahalanobis distance from mixed continuous and discrete data. Biometrics 56:394401. Berry AC, Berry RJ. 1967. Epigenetic variation in the human cranium. J Anat 101:361379. Blom DE, Hallgrmsson B, Keng L, Lozada MC, Buikstra JE. 1998. Tiwanaku colonization: bioarchaeological implications

for migration in the Moquegua Valley. World Archaeol 30: 238261. Brown MB. 1977. Algorithm AS 116. Appl Statist 26:343351. Brooks N, Drake N, McLaren S, White K. 2003. Studies in geography, environment and climate. In: Mattingly DJ, Daniels CM, Dore JN, Edwards D, Hawthorne J, editors. The archaeology of Fazzan. Volume1, Synthesis. London: Society for Lib yan Studies. p 3775. Buikstra JE. 1972. Techniques for coping with the age-regressive nature of non-metric traits. Am J Phys Anthropol 37:431432. Buikstra JE, Ubelaker DH. 1994. Standards for data collection from human skeletal remains. Fayetteville, AR: Arkansas Archaeological Survey Report Number 44. Camps G. 1974. Les civilizations prehistoriques de lAfrique du Nord et du Sahara. Paris: Doin. Camps G. 1982. Beginnings of pastoralism and cultivation in north-west Africa and the Sahara: origins of the Berbers. In: Clark D, editor. The Cambridge history of Africa, Volume1: From the earliest times to c. 500BC. Cambridge: Cambridge University Press. p 548623. Carson A. 2006. Maximum-likelihood variance components analysis of heritabilities of cranial nonmetric traits. Hum Biol 78:383402. Charles-Picard G, Picard C. 1968. The life and death of Carthage. New York: Taplinger Publishing Co. Cheverud JM, Buikstra JE. 1981a. Quantitative genetics of skeletal nonmetric traits in the rhesus macaques on Cayo Santiago. I. Single trait heritabilities. Am J Phys Anthropol 54:4349. Cheverud JM, Buikstra JE. 1981b. Quantitative genetics of skeletal nonmetric traits in the rhesus macaques on Cayo Santiago. II. Phenotypic, genetic, and environmental correlations between traits. Am J Phys Anthropol 54:5158. Cheverud JM, Buikstra JE. 1982. Quantitative genetics of skeletal nonmetric traits in the rhesus macaques of Cayo Santiago. III. Relative heritability of skeletal nonmetric and metric traits. Am J Phys Anthropol 59:151155. Christensen AF. 1998. Skeletal evidence for familial interments in the Valley of Oaxaca, Mexico. Homo 49:273288. Cremaschi M, Di Lernia S. 1999. Holocene climatic changes and cultural dynamics in the Libyan Sahara. Afr Archaeol Rev 16:211238. Daniels C. 1989. Excavation and eldwork amongst the Garamantes. Libyan Studies 20:4561. Davies WV. 2003. Sobeknakht of ElKab and the coming of Kush. Egypt Archaeol 23:36. Delia D. 1988. The population of Roman Alexandria. T Am Philol Assoc 118:275292. De Souza P, Houghton P. 1977. The mean measure of divergence and the use of non-metric data in the estimation of biological distances. J Archaeol Sci 4:163169. Destro-Bisol G, Donati F, Coia V, Boschi I, Verginelli F, Caglia A, Tofanelli S, Spedini G, Capelli C. 2004. Variation of female and male lineages in sub-Saharan populations: the importance of sociocultural factors. Mol Biol Evol 21:16731682. Di Lernia S. 2006. Building monuments, creating identity: cattle cult as a social response to rapid environmental changes in the Holocene Sahara. Quatern Int 151:5062. Dodo Y. 1974. Nonmetrical cranial traits in the Hokkaido Ainu and the northern Japanese of recent times. J Anthropol Soc Nip 82:3151. Donlon DA. 2000. The value of infracranial nonmetric variation in studies of modern Homo sapiens: an Australian focus. Am J Phys Anthropol 113:349368. Edwards DN. 2007. The archaeology of Sudan and Nubia. Annu Rev Anthropol 36:211228. Fattovich R. 1997. Northeastern African states. In: Vogel JO, editor. Encyclopedia of precolonial Africa. Archaeology, history, languages, cultures and environments. London: AltaMira Press. p 484489. Filer JM. 1992. Head injuries in Egypt and Nubia: a comparison of skulls from Giza and Kerma. J Egypt Archaeol 78:281285. Finnegan M, Cooprider K. 1978. Empirical comparison of distance equations using discrete traits. Am J Phys Anthropol 49:3946.

American Journal of Physical Anthropology

12

E. NIKITA ET AL.
Mattingly DJ. 2003a. Background to the archaeology of Fazzan. In: Mattingly DJ, Daniels CM, Dore JN, Edwards D, Hawthorne J. The archaeology of Fazzan. Volume1, Synthesis. London: Society for Libyan Studies. p 137. Mattingly DJ. 2003b. Synthesis of human activities in Fazzan. In: Mattingly DJ, Daniels CM, Dore JN, Edwards D, Hawthorne J, editors. The archaeology of Fazzan. Volume1, Syn thesis. London: Society for Libyan Studies. p 327375. Mattingly DJ. 2003c. Religious and funerary structures. In: Mattingly DJ, Daniels CM, Dore JN, Edwards D, Hawthorne J. The archaeology of Fazzan. Volume1, Synthesis. London: Society for Libyan Studies. p 177235. Mattingly DJ. 2006. The Garamantes: the rst Libyan state. In: Mattingly DJ, McLaren S, Savage E, al-Fasatwi Y, Gadgood K, editors. The Libyan desert. Natural resources and cultural heritage. London: Society for Libyan Studies. p 189204. Mauny R. 1978. Trans-Saharan contacts and the Iron Age in West Africa. In: Fage JD, editor. The Cambridge history of Africa, Volume 2: From c.500 B.C. to A.D. 1050. Cambridge: Cambridge University Press. p 266341. McGrath JW, Cheverud JM, Buikstra JE. 1984. Genetic correlations between sides and heritability of asymmetry for nonmetric traits in rhesus macaques. Am J Phys Anthropol 64: 401411. Mitchell P. 2005. African connections: archaeological perspectives on Africa and the wider world. Walnut Creek, CA: AltaMira Press. Nikita E. 2010. The Garamantes of Fazzan: bioarchaeological evaluation of desert-induced stress and Late Holocene human migrations through the Sahara. Ph.D. thesis, University of Cambridge, Cambridge. Nordstrom HA. 1972. Neolithic and A-Group sites. Stockholm: Astrom Editions. Ossenberg NS. 1969. Discontinuous morphological variation in the human cranium. Ph.D. thesis, University of Toronto, Toronto. Ossenberg NS. 1984. Within and between race distances in population studies based on discrete traits of the human skull. Am J Phys Anthropol 45:701716. Ossenberg NS, Dodo Y, Maeda Y, Kawakubo Y. 2006. Ethnogenesis and craniofacial change in Japan from the perspective of nonmetric traits. Anthropol Sci 114:99115. Ottoni C. 2007. Holocene human peopling of Libyan Sahara. Molecular analysis of maternal lineages in ancient and extant ` populations of Fezzan. Ph.D. thesis, Universita degli studi di Roma Tor Vergata, Rome. Petrie WMF. 1907. Gizeh and Rifeh. London: Adamant Media Corporation. Phillipson DW. 2005. African archaeology. Cambridge: Cambridge University Press. Pilbrow V. 2006. Lingual incisor traits in modern hominoids and an assessment of their utility for fossil hominoid taxonomy. Am J Phys Anthropol 129:323338. Prowse TL, Lovell NC. 1995. Biological continuity between the A- and C-Groups in Lower Nubia: evidence from cranial nonmetric traits. Int J Osteoarchaeol 5:103114. Prowse TL, Lovell NC. 1996. Concordance of cranial and dental morphological traits and evidence for endogamy in ancient Egypt. Am J Phys Anthropol 101:237246. Saunders SR, Popovich F. 1978. A family study of two skeletal variants: atlas bridging and clinoid bridging. Am J Phys Anthropol 49:193204. Schillaci MA, Irish JD, Wood CCE. 2009. Further analysis of the population history of ancient Egyptians. Am J Phys Anthropol 139:235243. Scott GR, Turner CG. 1997. The anthropology of modern teeth: dental morphology and its variation in recent human populations. Cambridge: Cambridge University Press. Seielstad MT, Minch E, Cavalli-Sforza LL. 1998. Genetic evidence for a higher female migration rate in humans. Nat Genet 20:278280 Sjvold T. 1973. The occurrence of minor non-metrical variants in the skeleton and their quantitative treatment for population comparisons. Homo 24:204233.

Freeman MF, Tukey JW. 1950. Transformations related to the angular and square root. Ann Math Stat 21:607611. Gatto M. 1997. Regional variations in the so-called A-group culture of lower Nubia. In: Barich B, Gatto M, editors. Dynamics of populations, movements and responses to climatic change in Africa. Rome: Bonsignori Editore. p 105111. Green RF, Suchey JM. 1976. The use of inverse sine transformations in the analysis of non-metric cranial data. Am J Phys Anthropol 45:6168. Green RF, Suchey JM, Gokhale DV. 1979. The statistical treatment of correlated bilateral traits in the analysis of cranial material. Am J Phys Anthropol 50:629634. Grewal MS. 1962. The rate of genetic divergence in the C57BL strain of mice. Genet Res 3:226237. Gruneberg H. 1952. Genetical studies on the skeleton of the Mouse IV. Quasi-continuous variations. J Genet 51:95114. Haas C. 1997. Alexandria in Late Antiquity: topography and social conict. Baltimore: The Johns Hopkins University Press. Hafsaas-Tsakos H. 2009. The kingdom of Kush: an African centre on the periphery of the Bronze Age world system. Nor Archaeol Rev 42:5070. Hanihara T, Ishida H. 2001. Frequency variations of discrete cranial traits in major human populations. I. Supernumerary ossicle variations. J Anat 198:689706. Hanihara T, Ishida H, Dodo Y. 2003. Characterization of biological diversity through analysis of discrete cranial traits. Am J Phys Anthropol 121:241251. Harris EF, Sjvold T. 2004. Calculation of Smiths mean measure of divergence for intergroup comparisons using nonmetric data. Dental Anthropol 17:8393. Harris EF. 2008. Statistical applications in dental anthropology. In: Irish JD, Nelson GC, editors. Technique and application in dental anthropology. Cambridge: Cambridge University Press. p 3567. Hauser G, De Stefano GF. 1989. Epigenetic variants of the human skull. Stuttgart: Schweizerbart. Irish JD. 1998. Ancestral dental traits in recent sub-Saharan Africans and the origins of modern humans. J Hum Evol 34:8198. Irish JD. 2006. Who were the ancient Egyptians? Dental afnities among Neolithic through post-dynastic peoples. Am J Phys Anthropol 129:529543. Irish JD. 2010. The mean measure of divergence: its utility in model-free and model-bound analyses relative to the Mahalanobis D2 distance for nonmetric traits. Am J Hum Biol 22:378395. Keita SOY. 1988. An analysis of crania from Tell-Duweir using multiple discriminant functions. Am J Phys Anthropol 75:375390. Kennedy GE. 1986. The relationship between auditory exostosis and cold water: a latitudinal analysis. Am J Phys Anthropol 71:401415. Konigsberg LW. 1987. Population genetic models for interpreting prehistoric intra-cemetery biological variation. Ph.D. thesis, Northwestern University, Chicago. Konigsberg LW. 1990. Analysis of prehistoric biological variation under a model of isolation by geographic and temporal distance. Hum Biol 62:4970. Konigsberg L, Buikstra J. 2006. Population structure analysis from prehistoric skeletal material. Am J Phys Anthropol Suppl 42:115. Lefevre-Witier P, Aireche H, Benabadji M, Darlu P, Melvin K, Sevin A, Crawford MH. 2006. Genetic structure of Algerian populations. Am J Phys Anthropol 18:492501. Liverani M. 2000. The Garamantes: a fresh approach. Libyan Studies 31:1728. Lloyd A. 2000. The Late period (664332BC). In: Shaw I, editor. The Oxford history of ancient Egypt. Oxford: Oxford University Press. p 369394. Mattingly DJ. 2000. Twelve thousand years of human adaptation in Fezzan (Libyan Sahara). In: Barker G, Gilbertson D, editors. The archaeology of drylands. Living at the margin. London: Routledge. p 160179.

American Journal of Physical Anthropology

SAHARA: BARRIER OR CORRIDOR?

Sjvold T. 1977. Non-metrical divergence between skeletal population: the theoretical foundation and biological importance of C.A.B. Smiths mean measure of divergence. Ossa 4:1133. Smith CAB. 1977. A note on genetic distance. Ann Hum. Genet 40:463479. Sutter RC, Mertz L. 2004. Nonmetric cranial trait variation and prehistoric biocultural change in the Azapa Valley, Chile. Am J Phys Anthropol 123:130145. Sutter RC, Verano JW. 2007. Biodistance analysis of the Moche sacricial victims from Huaca de la Luna plaza 3C: matrix method test of their origins. Am J Phys Anthropol 132:193206. Trigger BG. 1976. Kerma: the rise of an African civilization. Int J Afr Hist Stud 9:121. Tyrrell A. 2000. Skeletal non-metric traits and the assessment of inter-and intra-population diversity: past problems and future potential. In: Cox M, Mays S, editors. Human osteology in archaeology and forensic science. London: Greenwich Medical Media. p 289306. Williams B. 1997. Egypt and sub-Saharan Africa: their interaction. In: Vogel J, editor. Encyclopedia of precolonial Africa. Archaeology, history, languages, cultures and environments. London: AltaMira Press. p 465472.

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Wilson A. 2003. Irrigation technologies: forragas, wells and eld systems. In: Mattingly DJ, Daniels CM, Dore JN, Edwards D, Hawthorne J, editors. The archaeology of Fazzan. Volume 1, Synthesis. London: Society for Libyan Stud ies. p 235279. Winder SC. 1981. Infracranial non-metric variation: an assessment of its value for biological distance studies. Ph.D. thesis, University of Indiana. Wood ET, Stover DA, Ehret C, Destro-Bisol G, Spedini G, McLeod H, Louie L, Bamshad M, Strassmann BI, Soodyall H, Hammer MF. 2005. Contrasting patterns of Y chromosome and mtDNA variation in Africa: evidence for sex-biased demographic processes. Eur J Hum Genet 13:867876. Wooding S, Ostler Ch, Prasad BVR, Watkins WS, Sung S, Bamshad M, Jorde LB. 2004. Directional migration in the Hindu castes: inferences from mitochondrial, autosomal and Y-chromosomal data. Hum Genet 115:221229. Wysner GM. 1945. The Kabyle People. New York: Privately Printed. Zakrzewski SR. 2003. Variation in ancient Egyptian stature and body proportions. Am J Phys Anthropol 121:219299.

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