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DNA ANALYSIS OF MOCHE AND SICAN POPULATIONS:
RESULTS AND IMPLICATIONS
Ken-ichi Shinoda Department of Anatomy Saga Medical School, Saga, Japan
Izumi Shimada Department of AnthropologySouthern Illinois University, Carbondale, IL 62901
Walter Alva Museo Arqueológico Brüning Lambayeque, Perú
Santiago Uceda Museo Arqueológico Universidad Nacional de Trujillo, Trujillo, Perú
Evolving out of the earlier interdisciplinary investigation of Middle Sicán (ca. A.D. 1000) burials and mortuary practices by the Sicán Archaeological Project (SAP), Shinoda and Shimada have been involved in the first, large-scale mitochondrial DNA (mtDNA) analysis of prehispanic Moche (a.k.a. Mochica) and succeeding Sicán (a.k.a. Lambayeque) populations on the north coast of Peru. The present study discusses results of the analysis of human remains of diverse time periods, social status and roles, and, perhaps, ethnic origins excavated over the past 15 years at the sites of Sipán in the mid-Lambayeque Valley and Moche (Temple of the Moon/Huaca de la Luna) in the lower Moche Valley. In addition, mtDNA data of burials excavated by the SAP at the Middle Sicán capital of Sicán are presented for genetic comparison between the sampled Moche and Sicán populations as well as defining their respective internal variabilities. Research questions that guided the present study include: Were the Moche social elite of different valleys biologically related or unrelated? Who were the sacrificial victims discovered at the Huaca de la Luna vis-a-vis the "warriors" and others who had been carefully interred in and around the temple? Were the Moche elite personages from Sipán related, together constituting a regional dynasty? Were the Moche elite at Sipán related to the later Sicán elite? Genetic material recovered from a single tooth from each of the sampled individuals enabled us to establish their mtDNA sequences and haplotypes. Maternally related individuals were identified and haplotype comparison of these individuals with others both supports and questions existing views concerning Moche and Sicán societie.
RESEARCH ISSUES AND SETTINGS AND BURIAL SAMPLES
In 1990 the Sicán Archaeological Project (SAP) initiated its long-term investigation into social, ideological, and demographic dimensions of the Middle Sicán culture (A.D. 900-1100) through interdisciplinary analysis of its burials and mortuary practices. Based on burial pattern and locations elucidated through prior research, Middle Sicán burials representing different segments of the social spectrum were excavated in different areas within the capital site of Sicán in the mid-La Leche Valley (Poma National Historical Sanctuary; Fig. 1). The burials included 5 and 24 individuals found inside two deep shaft tombs at the north base of the major adobe platform mound of Huaca Loro called the "East" and "West" tombs, respectively (Figs. 2, 3). Togain a comprehensive understanding of the biological and social relationships among individuals within and between excavated tombs, systematic analyses were conducted of inherited dental traits, mtDNA, a variety of health-dietary indicators, and cranial deformation. To complement these analyses, we also conducted stylistic, distributional, compositional, and materials analyses of associated grave goods.
Individually and collectively, these analyses revealed important biological and social relationships among the excavated Middle Sicán burials from Huaca Loro (e.g., Corruccini and Shimada 2002; Farnum and Shimada 1999; Farnum et al. 2001a, b; Shimada et al. 1998, 2000, 2001, n.d.). Our interest in gaining an understanding of the broader significance of the mtDNA data from Huaca Loro has led us to seek comparative samples, including earlier Moche burials from Sipán and burials and sacrificial victims from Moche.
At an adobe plaform (Mound 1) within the extensive, multi-component site of Sipán in the mid-Lambayeque Valley (Fig. 1, 4), some 42 km south of the site of Sicán, a team from the Bruning Regional Archaeological Museum led by Walter Alva has conducted an exhaustive excavation of burials following the discovery of looting of a major Moche tomb in 1986. Among others, they have unearthed 12 early Moche tombs of different eras (first to fourth centuries A.D.) and social rank (e.g., Alva 1994, 2001; Alva and Donnan 1993). These burials together are believed to represent "four broad categories: ruling lords, priests, military leaders, and the second-rank assistants to military leaders and priests" (Alva 2001:223). Those related to political or military power were interred in the northern section of the funerary platform, and religious personnel in the southern part (Alva 2001:223).
Tombs 1 and 3, representing the most elaborate burials, pertain to the "ethnic lords" commonly referred to as the "Lord of Sipán" and "Old Lord of Sipán", respectively . Tomb 1 (5 x 5 m; pertaining to the last of the six-phase construction sequence [Phase 6]) excavated in 1987 contained two adult males (35-50 years old), three females (ca. 15-20 years old) and a child (ca. 9-10 years) placed in cane coffins around the central wooden coffin (Fig. 5). Inside the central coffin was the Lord of Sipán, an adult male of 35-45 years of age. The fact that all the female skeletons were disarticulated and jumbled suggest that they were secondary burials. Above the coffin were an adult male "guard" (ca. 20 year old), whose feet had been cut off and a seated adult male, presumably a retainer.
Tomb 3, a simple pit (Phase 1) placed near the bottom of Mound 1, contained the Old Lord, an adult male of ca. 45-55 years of age. He was accompanied by a young adult female (16-18 years old). Alva (2001:229) believes that he was "perhaps some four or five generations before the Lord of Sipán in Tomb 1."
Tomb 2 (Phase 6) that was placed in the final construction phase like Tomb 1 contained a wooden coffin with the body of the Priest, an adult male of ca. 35-45 years old (Fig. 6). As with the Lord, the Priest was accompanied by a young male (ca. 14-17 years old), two young women (18-20 years old and 19-25 years old), and one juvenile (8-10 years old), each in his or her own cane coffin. In addition, there was a young adult male "guard" with his feet amputated placed above the burial chamber, much like the arragement seen in Tomb 1.
The remaining six excavated tombs are less sumptuous than those described above and believed to be elite individuals of "lesser rank" who may have served as assistants of warriors and priests.
Based on the patterned distribution in time and space of the 12 tombs, functional interpretations of associated grave goods, and the variation in the composition and quality of these goods, the excavated individuals have been interpreted as members of the regional Moche royal family and their attendants. Though the spatial grouping, particularly that found within each tomb, is highly suggestive of social and biological relationships among the excavated individuals, inferences based on the material dimension of the mortuary treatments remained inconclusive. For example, based on observed similarities in type and style of associated personal ornaments, the Old Lord is believed to have been an ancestor of the Lord four to five generations earlier. Can we provide independent support for this inference? If Mound 1 served as a royal mausoleum, what other familial ties existed among the excavated individuals? Also, given their relative proximity to each other and the cross-culturally common practice of marriage alliances among social elite, could the Moche and later Sicán elite individuals have been related to each other?
Huaca de la Luna, Moche
Between 1995 and 1998, to test his hypothesis concerning Moche human sacrificial rituals, Steve Bourget (1997a,b,c, 2001) excavated enclosed Patio 3A (50 x 10-17 m) and the adjacent, conterminous construction of Platform II (50 x 17 m) toward the rear of the famed Huaca de la Luna at the site of Moche in the lower Moche Valley, some 175 km southeast of Sipán (Figs. 1, 7). His excavation of the Patio revealed 15 strata of human remains representing at least six distinct rituals and 70 young adult males (Fig. 8; Bourget 1997a,b,c, 2001; Bourget and Newman 1998; Verano 1998, 2001). Decapitated, dismembered or otherwise mutilated bodies and parts were strewn around a rock outcrop, the inferred natural sacrificial altar, embedded in dried mud washed in by torrential rains and subsequently covered by aeolian sand. The head injuries found on many crania seemed to have been caused by hard blows with war maces. Evidence clearly points to gruesome human sacrificial rituals carried out in front of the outcrop during various episodes of El Niño-related rains that date to around the end of the sixth century A.D. (Moche Phase IV).
Excavation of the adjacent Platform II revealed the partially looted burials of two old men (Tombs 1 and 3) who are thought to have been "warrior-priests" involved in the capture of sacrificial victims and/or their execution in the adjacent Patio (Bourget 2001; Bourget and Newman 1998). The associated ceramics and sculptures showed representations of death, warfare, prisoners and sacrifice. In fact, Tomb 1 contained a wooden mace covered with a thick layer of human blood.
Excavations near the base of the Temple mound yielded a series of Moche burials (Phases III and IV) of relatively high social status, including two burials of inferred potters who were accompanied by diverse grave goods (e.g., Chapedalaine 1997, 2001; Uceda 1997; Uceda and Armas 1998; Uceda and Mujica 1997). The "potters" are believed to have been social elite who had “some control over the Moche ideological and cosmological realm through pottery production" (Uceda and Armas 1998:107).
The two contrasting groups of excavated individuals – those who were carefully interred and those who were violently killed – offered a unique opportunity to shed light on long-standing debate over the identity of sacrificial victims.
For decades, much has been written about the numerous realistic drawings, murals, and sculptural representations of “warriors” and armed combat in Moche art. Interpretations range from the one that argues for intertribal conflicts resulting from the lack of a strong authority to dominate and unite all the Moche groups in the different valleys (Kutscher 1955: 28, 1967:120) to the one which sees representations as showing ritualistic battles fought between young men of rival moities of the Moche society (Hocquenghem 1978, 1987). Most commonly, archaeologists (e.g., Conklin and Moseley 1988; Dillehay 2001; Topic 1982; Wilson 1987, 1988) have seen the combat as a reflection of the importance of warfare as the means of Moche political dominance over other coexisting or nearby ethnic groups such as the Gallinazo and Recuay and consequent territorial expansions. On the other hand, systematic studies of these images indicate the primary function of combat was to capture live “warriors” for subsequent sacrifice (e.g., Bourget 2001; Donnan 1997; Hocquenghem 1987; Shimada 1994). But, who was fighting? Moche versus non-Moche (ethnically speaking) or competing groups within Moche society? Were sacrificial rituals the cause or effect of the combat?
These divergent interpretations carry with them different implications testable through genetic analysis of "warriors" and their sacrificial victims (defeated "warriors"). It is the recognition of this fact that led us to apply mtDNA analysis. More specifically, we were interested in defining the genetic profiles of and linkages, if any, (1) between the sacrificial victims, on the one hand, and "warriors" and other members of the “victorious” group, on the other, excavated at the site of Moche, and (2) between the social elite of the sites of Moche and Sipán. The latter aim relates to the question of the sociopolitical composition and organization of what the archaeologist calls the Moche or Mochica culture.
There are additional burials being analyzed. For example, those from Huaca Las Ventanas and Rodillona at Sicán and Huaca Sialupe in lower Leche Valley are expected to provide a better picture of intra-regional mtDNA variability during the Middle Sicán period. On the other hand, those from the Huaca Cao Viejo within the El Brujo Complex near the mouth of the Chicama Valley include elite and commoner individuals of the Moche (Phases III and IV) and Sicán periods, allowing us to explore the genetic variation in time and space as well as across social boundaries. Reults of their analysis will be reported in near future.
There are various methods available for exploring kinship relationships among excavated burials. Biomolecular methods based on analysis of ancient DNA offer vastly superior discriminatory power over other methods, but the preservational problem also looms large. Ancient nuclear DNA (nDNA) degrades much more easily than mtDNA and thus its analysis cannot be applied to most excavated skeletons (e.g., Alcamo 2001; Brown and Brown 1992).
Human mtDNA is a circular genome of approximately 16500 base pairs in length (Fig. 9). The complete nucleotide sequences have been reported. Thus we could amplify the DNA of the desired region by Polymerase Chain Reaction (PCR). MtDNA has been widely used as a tool in the study of human evolution because it is maternally inherited and nonrecombining. Moreover, mtDNA accumulates base changes at a rate 5-10 times that of nDNA, and the D-loop region is its fastest-changing segment; therefore, substantial sequence variations exist among individuals. DNA extracted from ancient materials is considered to be in low concentration and degraded condition. A major advantage of mtDNA is that it is present in many copies, up to perhaps a thousand per cell. Thus, it has a greater likelihood of being amplified than nDNA, which occurs in only one copy per cell. Often a sufficient amount of mtDNA can be extracted from a single tooth, an important consideration given the destructive nature of the analysis.
It is for the above reasons that we adopted mtDNA analysis for our research. At the same time, the analysis requires tremendous rigor and time largely due to the less than ideal condition of the samples and their potential contamination. Further, this analysis can only establish genetic linkages through maternal lines as mtDNA genes are not re-arranged nor is its DNA subdivided during reproduction unlike nDNA (Alcamo 2001:226).
Tooth samples were used in our study of Sicán burials. It has been argued that teeth form an effective natural barrier to exogenous DNA contamination and that the DNA from teeth appear to lack most of the inhibitors to enzymatic amplification of ancient DNA (Woodward et al. 1994). A well-preserved tooth was extracted from each individual by Shinoda in Peru. In our study, a combination of RFLP haplotype and D-loop sequence analysis was used to characterize mtDNA variation. Extracted DNA was amplified by PCR. When the amount amplified was not sufficient, the second PCR was performed for the next direct sequencing step in D-loop amplification. In case several ambiguous positions were observed in the sequencing data, possibly due to mismatch by the DNA polymerase during PCR, the next DNA cloning step was performed. Analytical details are described elsewhere (Shimada et al. n.d.). The suspected false positive results stemming from contamination with contemporary DNA (Lawlor et al. 1991) and other questionable data (e.g., Kolman and Tuross 2000) were excluded. Individuals whose base sequences in the D-loop region of the mtDNA and attendant haplotypes have been determined are presented in Table 1 and Figure 9.
RESULTS AND THEIR INTERPRETATION
Our mtDNA analysis has revealed some significant biological relationships as well as haplotype patterning that merit our close examination.
Among the Middle Sicán burials from Huaca Loro at Sicán, kinship ties were defined among 12 out 22 relatively young women who were divided into two groups that are symmetrically opposed (north-south) in respect to the centrally placed principal burial of the tomb (Figs 10, 11). As seen in Table 1, six individuals in the West Tomb had unique haplotypes, and the remaining individuals could be classified into four haplotype groups. The spatial distribution of the defined haplotypes clearly cluster into two groups, i.e., those identified by triangles and diamonds on the north side as opposed to squares and open circles on the south side in Figure 11. In other words, the north vs. south groupings of women has a kinship basis. The south group women who show a relatively high degree of genetic affinity among them in terms of mtDNA and bilaterally inherited dental traits may well represent an endogamous group (see Corruccini and Shimada 2002). The north group situation is not as clear. Independent lines of evidence corroborating this conclusion from analyses of inherited dental traits, nutritional-health indicators, and associated artifacts are presented elsewhere (e.g., Corruccini and Shimada 2002; Farnum and Shimada 1999; Farnum et al. 2001; Shimada 1995; Shimada et al. 1998, 2000, 2001, n.d.)
That these two groups may also represent members of two distinct ethnic groups is suggested by a notable difference in the style of ceramics that accompanied them. The north side women are predominantly associated with vessels that show a strong affinity to the earlier Moche style, while the south side women are found with typical Middle Sicán style vessels.
Comparison of the haplotype composition of these two groups with that of the Sipan burials and modern populations farther afield also lends additional support.
Among 16 early Moche individuals from Sipán we sampled, ten yielded sufficient genetic materials to determine their mtDNA sequences (Table 1). Seven mtDNA types were identified among them. Though their sample sizes are small, comparison of Sicán and Sipán samples suggest they may well have two distinct genetic background. Haplogroups C and D predominate the Sipán individuals, while haplogroups A, B, and other newly defined groups characterize the Sicán individuals from Huaca Loro. In fact, in terms of the haplogroup variability and ratios, the Sipán sample is similar to the modern Central Andean populations, while the Sicán sample shows a clear affinity to the North Andean populations (Please insert relevant references) . Overall, what emerges is the distinct possibility that the north and south side women represent members of the two distinct ethnic groups integrated under the Middle Sicán polity.
MtDNA analysis of Moche burials from Sipán (Table __) provided important independent support to the notion that Mound I at Sipán was in essence a mausoleum for members of a regional elite family and close associates. The analysis showed that the Lord, the Old Lord, a young woman and a young man who accompanied the Priest were all maternally related to each other. A straightfoward interpretation of the relationship between the Old Lord and the Lord would be that they were either two brothers or cousins or an uncle and a nephew. Given the significant stratigraphic difference between their tombs, they are more likely to be an uncle-nephew pair several generations removed.
Two distinct generations of male rulers who are maternally related could result in a number of ways. Based on ethnohistorical documents pertaining to northern coastal region (yunga language groups), Rostworowski (2000:178) described the prevalent rule of succession for theyunga nobles, "in which it was the brother of the deceased rather than his son who assumed power" and that this "process continued through to the last brother of that generatioin, before passing to the generation below." Another possibility is the "avunculate" system as practiced by the Inkas "in which the son of the sovereign's sister had priority over the son of the sovereign", giving rise to "cases of incestuous marriage between ruling brother and sister . . . ." (Rostworowski 2000:178). Though these scenarios are suggestive , it should be kept in mind that any attempt to equate the theoretical rules and the actual succession is difficult since the latter could significantly diverge from the former for a variety of reasons.
The documented biological relationship between the Priest and two of his four accompanying individuals also brings up a similar range of possibilities. For example, we may think of their relationship as that of an uncle and his favorite nephews who served him in life. Conversely, the Priest and the woman may be seen as two siblings or cousins who married each other. The latter scenario is weakened by the fact that both there were no corresponding relationships between the Lord and the Old Lord, on the one hand, and their female companion(s), on the other. If the origin myth of the protohistorical Chimú Kingdom of the North Coast that explains how the social elite and commoners were derived from distinct stars (e.g., Rowe 1948) is any reflection of the earlier Moche social reality, then, endogamy or even sibling or cousin marriage among the Moche elite members in Lambayeque remains a strong possibility. In this regard, it is notable that there is no linkage along maternal line between individuals of the highest social "rank" and their associates, on the one hand, and those of the second "rank" and their inferred "retainers", on the other.
What is notable about the Huaca de la Luna data is the exceptional haplotype homogeneity (Table __); that both the sacrificial victims and other interred indivdiuals share the same haplotype, though no maternal kinsmen were identified. It is quite likely that these two groups pertain to a single population that experienced a genetic bottleneck (founder's effect) in a relatively recent past. For example, the warriors and other individuals of relatively high social status at the site of Moche may have been descendants of a small homogenous group that colonized the area or alternatively suffered a drastic reduction in number (due to warfares or epidemics). Given that relatively small samples of Moche burials at Sipán and El Brujo show much more haplotype variabilities, we must consider the distinct possibilities that the individuals sacrified at Huaca de la Luna pertained to the same local population as the sacrificers and other residents of the site of Moche. In other words, we need to consider the possibility that at least in some sacrificial rituals, victims may have been "caught" as result of intra-group ritual combats. It can be further argued that Moche elite groups of the mid-Lambayeque, lower Chicama and lower Moche valleys are likely to represent distinct local breeding groups, lending some support to a view that what we call the Moche polity may have been in reality an aggregation of a series of regional polities (Castillo and Donnan 1994; Shimada 1994). The possibility that the sacrificial victims pertained to what archaeologists call Cajamarca and Recuay societies centered in the adjacent upper valleys and highlands seems remote.
Though tantalizing, these conclusions must be regarded as tentative as they are based on small samples and inadequate understanding of the genetic processes and variabilites of pertinent regional populations.
The D-loop region sequences of this study, encompassing the 154 nucleotides between mtDNA positions 16209 and 16362 were compared with the previously published sequences of 111 Amerindians of Central and South America (Kuna-Ngobe, Batista and Kolman 1995, Kolman et al. 1995; Panama, Santos et al. 1994; Cayapa, Rickalds et al. 1999; Amazon, Ward et al. 1996; Brazil, Alves-Silva et al. 2000; and South American mummies, Monsalve et al. 1996). The haplogroups of these samples were defined on the basis of RFLP analysis. The relationship among these populations was examined by using the median network method (Bandelt et al. 1994). Open circles represent distinct mtDNA types found in the present study and Amerindians, and black circles represent types that are absent from those populations. Four clusters of Native Americans previously identified by other authors were classified. Colored circles represent mtDNA types seen in the present study. Reticulations in the network indicate the existence of incompatible nucleotide configurations. Because the pattern of nucleotide substitution in this region is quite complicated, it is difficult to reconstruct the precise tree from such a short nucleotide sequence. Therefore, the haplogroup status can be read off only to some extent. There is room for further investigation to decide the precise relationships among these populations. Illustration by K. Shinoda.
Our study effectively demonstrated the value of the mtDNA analysis of well-preserved prehispanic human burials in elucidating their biological and social relationships. For the well-preserved burials from coherent contexts (e.g., intra-cemetery samples) from Sipán and Sicán, the analysis revealed specific maternal kinship ties among some of the burials, providing important corroborative evidence for the exisiting archaeological interpretations on the organizational principles and other aspects of the excavated burials. However, without an adequate baseline knowledge of demographic processes that operated in the past and the overall genetic variability of each regional population, exploration into biological relationships among samples derived from different valleys is inconclusive and tenuous. At the same time, the genetic homogeneity revealed for the Huaca de la Luna sample is so striking that it leads us to suggest that the sample represents a population that went through a genetic bottleneck and is probably distinct from the Moche population to which Sipán individuals belonged. It is also likely that both the sacrificers and sacrificial victims both pertained to the same ethnic group as opposed to the Recuay or Cajamarca groups. We end this paper with a note of caution: It should be kept in mind that mtDNA data do not inform us of any genetic connections along the paternal line. To specify biological relationships among excavated individuals beyond what are described in this paper is inadvisable without additional independent lines of evidence.
We are grateful to Steve Bourget, members of the Bruning Regional Archaeological Museum, Proyecto Huaca de la Luna, and Sicán Archaeological Project, and Yutaka Yoshii for their assistance in collection of tooth samples used in the mtDNA analysis. Research by Shinoda for this study was supported by a grant from the Ministry of Education of Japan.
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