Beruflich Dokumente
Kultur Dokumente
Manchester Interdisciplinary Biocentre, Faculty of Life Sciences, University of Manchester, 131 Princess Street, Manchester M1 7DN, UK
The Manchester Museum, University of Manchester, Oxford Road, Manchester M13 9PL, UK
a r t i c l e i n f o
a b s t r a c t
Article history:
Received 28 January 2008
Received in revised form 7 April 2008
Accepted 21 April 2008
The richness of the burials in Grave Circle B at Mycenae, Greece indicates that the 35 people interred
there held elite status during their lifetimes 3500 years ago. It has been speculated that the groups of
burials represent different dynasties or branches of the same family. To test this hypothesis, we carried
out an exhaustive ancient DNA (aDNA) study of 22 of the skeletons. We were unable to identify nuclear
aDNA in any specimen, but we obtained authentic mitochondrial aDNA sequences for four individuals.
The results were compared with facial reconstructions and interpreted within the archaeological context
represented by the organisation of the graves and the positions of the burials within the graves. We
conclude that the contemporaneous male G55 and female G58 skeletons, which both possess the UK
mitochondrial haplogroup, were brother and sister. The implication is that G58 was buried in Grave
Circle B not because of a marital connection but because she held a position of authority by right of birth.
The results illustrate the difculty in using aDNA to study kinship relationships between archaeological
specimens, but also show that aDNA can advance understanding of kinship when used to test hypotheses
constructed from other evidence.
2008 Elsevier Ltd. All rights reserved.
Keywords:
Ancient DNA
Kinship analysis
Mitochondrial DNA
Mycenae
Polymerase chain reaction
1. Introduction
The Bronze Age citadel at Mycenae in Greece is one of the most
evocative prehistoric sites in all of Europe. The legendary home of
Agamemnon and Clytemnestra, Mycenae held a natural attraction
for early antiquarians in the years before its rst systematic study
by Heinrich Schliemann in the 1870s. Schliemanns famous telegram, sent during his excavation of Grave Circle A in 1876, stating
that he had gazed upon the face of Agamemnon, turned out to be
erroneous for the burials that he had uncovered predated the
Trojan War by some four centuries, but his excavations were
nonetheless signicant as they established Mycenae as one of the
richest and, by implication, most powerful of the Aegean states
during the 17th to 12th centuries BC.
The burials in Grave Circles A and B span some ve generations
during the period 16501500 BC, Grave Circle B predating A with
possibly 50 years overlap between the two. The Grave Circles
therefore date to the very beginning of the Mycenaean age, the
cusp of the Middle to Late Helladic periods when Mycenae was
establishing itself as a dominant trading and political power in
the eastern Mediterranean. Within Grave Circle B there is a
development from simple cist burials to larger, deeper and richer
* Corresponding author. Tel.: 44 161 306 4173; fax: 44 161 306 5201.
E-mail address: terry.brown@manchester.ac.uk (T.A. Brown).
0305-4403/$ see front matter 2008 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jas.2008.04.010
Shaft Graves, while Grave Circle A comprises six Shaft Graves. Many
of the burials in Grave Circle B were accompanied by weapons,
pottery and/or gold ornaments; one, in Grave G, was buried with
a face-mask made of electrum (a naturally occurring goldsilver
amalgam). Generally they were less well endowed than the remarkable gold-laden burials in Circle A, but the richness of both
Grave Circles leaves little doubt that their occupants were elite
members of early Mycenaean society. These considerations prompt
questions about the relationships within this elite group, in particular whether the individuals were members of a single family or
small number of families who had established themselves as the
ruling dynasty in early Mycenae. Schliemann believed that the gold
face-masks of Grave Circle A were portraits of the owners and noted
their dissimilarities (Schliemann, 1876), but this observation did
not hinder speculations that the 19 individuals buried in Circle A
were members of a single family (Mylonas, 1957). When Grave
Circle B was discovered in 1951 this assumption gained weight as
the graves in this cemetery appear to be laid out in four groups,
each group spanning the different phases of the site, reminiscent of
family plots. Dr J.L. Angel, who examined the Circle B skeletons in
1954, felt he could visualise the faces that lay over the skulls, and
suggested family relationships (Angel, 1973). Forty years later,
modern techniques of facial reconstruction were applied to the
seven best preserved skulls, the results suggesting that these seven
individuals fall into three groups, the heart-shapes comprising
G55, G58 and A62, the long faces of G51 and Z59, and the beaky
face of B52, with S131, the earliest of these seven burials, having
features in common with both of the rst two types (Musgrave
et al., 1995; Prag and Neave, 1997) (Fig. 1).
Ancient DNA (aDNA) analysis has the potential to identify kinship patterns between groups of skeletons, maternal relationships
being revealed by mitochondrial DNA (mtDNA) typing, paternal
ones by studying markers such as short tandem repeats (STRs) on
the Y chromosome, and general family relationships by typing
autosomal STRs. Realisation of this potential has, however, been
hampered by the problems inherent in aDNA research and
although success has been achieved with historic material (Gill
et al., 1994; Gerstenberger et al., 1999; Dudar et al., 2003; Gilbert
et al., 2007) there have been few authenticated kinship studies with
archaeological specimens. Here we describe an exhaustive aDNA
study of the Grave Circle B skeletons which has resulted in
assignment of mtDNA haplogroups to four skeletons, providing
insights into the family relationships between these individuals.
2581
2582
Table 1
Results of PCRs
Skeleton
G51
B52
P53
H54
G55
L56
X57
G58
Z59
D60
D61
Extractions
MtA
MtG
MtC
MtF
MtD
MtW
MtV
MtZ
CD4
D1S656
D2S1338
D2S1358
D5S818
D6S366
D8S535
D8S1179
D10S1248
D10S2325
D14S1434
D16S539
D18S51
D22S1045
FGA
TH01 A
TH01 B
VWA
DYS389
DYS391
DYS393
DYS426
DYS460
M35
M173
GA
MB
2
2/0/0
3/0/0
8/3/0
4/2/0
6/4/0
1/1/0
3/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
4/0/0
1/0/0
1
1/0/0
1/0/0
2/0/0
1/1/0
1/0/0
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/1/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/1/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2
2/0/0
3/2/2
8/6/3
4/3/2
6/4/1
1/1/0
3/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/1/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/1/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
4/1/1
1/0/0
1
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/1/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/1/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1
1/0/0
1/0/0
2/0/0
1/1/0
1/0/0
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
2
2/0/0
3/2/2
8/6/3
4/3/2
6/4/1
1/1/0
3/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
4/0/0
1/0/0
2
2/0/0
3/0/0
7/5/2
4/2/2
6/4/1
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
3/0/0
1/0/0
1
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
Skeleton
A62
Q63
N66
N66a
I68
A69
K70
L70a
L70a1
L70a2
L70a3
Extractions
MtA
MtG
MtC
MtF
MtD
MtW
MtV
MtZ
CD4
D1S656
D2S1338
D2S1358
D5S818
D6S366
D8S535
D8S1179
D10S1248
D10S2325
D14S1434
D16S539
D18S51
D22S1045
FGA
TH01 A
TH01 B
VWA
DYS389
DYS391
DYS393
DYS426
DYS460
M35
M173
GA
MB
2
2/0/0
3/2/2
8/6/2
4/3/2
6/4/0
1/1/0
3/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
4/0/0
1/0/0
1
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1
1/0/0
1/0/0
2/1/0
1/1/0
1/0/0
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1
1/0/0
1/0/0
1/0/0
1/1/0
1/0/0
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1
1/0/0
1/0/0
2/0/0
1/1/0
1/0/0
1/1/0
2/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/1/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
1/0/0
1/0/0
2/0/0
1/0/0
1/0/0
2/0/0
1/0/0
Key: Number of PCRs/Number giving a band of the correct size/Number of bands thought to result from endogenous DNA.
2583
2584
References
Angel, J.L., 1973. Human skeletons from grave circles at Mycenae. In: Mylonas, G.E.
(Ed.), Ho Taphikos Kyklos B ton Mykenon, vol I. Athens Archaeological Society,
Athens, pp. 379397.
Bouwman, A.S., Brown, T.A., 2005. The limits of biomolecular palaeopathology:
ancient DNA cannot be used to study venereal syphilis. J. Archaeol. Sci. 32, 691
702.
Bouwman, A.S., Chilvers, E.R., Brown, K.A., Brown, T.A., 2006. Identication of the
authentic ancient DNA sequence in a human bone contaminated with modern
DNA. Am. J. Phys. Anthropol. 131, 428431.
Brown, T.A., Brown, K.A., Flaherty, C.F., Little, L.M., Prag, A.J.N.W., 2000. DNA analysis
of bones from Grave Circle B at Mycenae: a rst report. Annu. Brit. Sch. Athens
95, 115119.
Butler, J.M., Chen, Y., McCord, B.R., 2003. The development of reduced size STR
amplicons as tools for analysis of degraded DNA. J. Forensic Sci. 48, 10541064.
Chilvers, E.R., 2004. Ancient DNA and palaeopathology: malaria and Ancient Greece.
PhD Thesis, University of Manchester Institute of Science and Technology.
Chilvers, E.R., Bouwman, A.S., Brown, K.A., Brown, T.A. Ancient DNA in human bones
from Neolithic and Bronze Age sites in Greece and Crete. J. Archaeol. Sci., in
press.
Coble, M.D., Butler, J.M., 2005. Characterization of new miniSTR loci to aid analysis
of degraded DNA. J. Forensic Sci. 50, 4353.
Cooper, A., Poinar, H., 2000. Ancient DNA: do it right or not at all. Science 289, 1139.
Dudar, J.C., Waye, J.S., Saunders, S.R., 2003. Determination of a kinship system using
ancient DNA, mortuary practice, and historic records in an Upper Canadian
pioneer cemetery. Int. J. Osteoarchaeol. 13, 232246.
Evison, M.P., 2001. Ancient DNA in Greece: problems and prospects. J. Radioanal.
Nucl. Chem. 247, 673678.
Gerstenberger, J., Hummel, S., Schultes, T., Hack, B., Herrmann, B., 1999.
Reconstruction of a historical genealogy by means of STR analysis and
Y-haplotyping of ancient DNA. Eur. J. Hum. Genet. 7, 469477.
Gilbert, M.T.P., Bandelt, H.-J., Hofreiter, M., Barnes, I., 2005. Assessing ancient DNA
studies. Trends Ecol. Evol. 20, 541544.
Gilbert, M.T.P., Djurhuus, D., Melchior, L., Lynnerup, N., Worobey, M., Wilson, A.S.,
Andreasen, C., Dissing, J., 2007. MtDNA from hair and nail claries the genetic
relationship of the 15th century Qilakitsoq Inuit mummies. Am. J. Phys.
Anthropol. 133, 847853.
Gill, P., Ivanov, P.L., Kimpton, C., Piercy, R., Benson, N., Tully, G., Evett, I., Hagelberg, E.,
Sullivan, K., 1994. Identication of the remains of the Romanov family by DNA
analysis. Nat. Genet. 6, 130135.
Musgrave, J.H., Prag, A.J.N.W., Neave, R.A.H., Thimme, D.L., 1995. Seven faces from
Grave Circle B at Mycenae. Annu. Brit. Sch. Athens 90, 89100.
Mylonas, G.E., 1957. Ancient Mycenae: the Capital City of Agamemnon. Routledge &
Kegan Paul, London.
Prag, A.J.N.W., Neave, R.A.H., 1997. Making Faces. British Museum Press, London
(Chapter 6).
Pruvost, M., Schwarz, R., Bessa Correia, V., Champlot, S., Braguier, S., Morel, N.,
Fernandez-Jalvo, Y., Grange, T., Geigl, E.-M., 2007. Freshly excavated fossil bones
are best for amplication of ancient DNA. Proc. Natl. Acad. Sci. U.S.A. 104, 739
744.
Richards, M., Macaulay, V., Hickey, E., Vega, E., Sykes, B., Guida, V., Rengo, C.,
Sellitto, D., Cruciani, F., Kivisild, T., Villems, R., Thomas, M., Rychkov, S.,
Rychkov, O., Rychkov, Y., Golge, M., Dimitrov, D., Hill, E., Bradley, D., Romano, V.,
Cali, F., Vona, G., Demaine, A., Papiha, S., Triantaphyllidis, C., Stefanescu, G.,
Hatina, J., Belledi, M., Di Rienzo, A., Novelletto, A., Oppenheim, A., Norby, S., AlZaheri, N., Santachiara-Benerecetti, S., Scozzari, R., Torroni, A., Bandelt, H.-J.,
2000. Tracing European founder lineages in the Near Eastern mitochondrial
gene pool. Am. J. Hum. Genet. 67, 12511276.
Schliemann, H., 1876. Letter to The Times dated 15 November 1876.
Smith, C.I., Chamberlain, A.T., Riley, M.S., Stringer, C., Collins, M.J., 2003. The thermal
history of human fossils and the likelihood of successful DNA amplication.
J. Hum. Evol. 45, 203217.