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Tutankhamen's family
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PostPosted: Wed Feb 17, 2010 2:21 am    Post subject: Reply with quote

In addition to the STEVOR and AMA1 genes, we attempted amplification of alleles of the MSP1 and MSP2 genes specific to P falciparum. Because of the fragmented nature of the ancient DNA, we did not obtain positive amplifications when targeting the larger (>400 bp) PCR alleles of the MSP2 gene but were successful in amplifying different alleles of the MSP1 gene (for details on MSP1 data, see eAppendix).29-30 Using extracts from Tutankhamun and Yuya, we repeatedly amplified the RO33 and MAD20 alleles, which is indicative of at least a double infection with the P falciparum parasite. The DNA of Thuya yielded amplicons for the RO33 allele. The DNA of TT320-CCG61065 was refractory to MSP1 amplifications. Cloning the obtained allelic fragments into TA plasmid vectors and subsequent Sanger sequencing of 21 clones designated the sequences as specific for MSP1 (eAppendix).
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PostPosted: Wed Feb 17, 2010 2:23 am    Post subject: Reply with quote

COMMENT

Kinship Determination

More than 55 bone biopsies were used to elucidate the individual relationships of 18th-dynasty individuals, with the result that several of the anonymous mummies or those with suspected identities are now able to be addressed by name. These include KV35EL, who is Tiye, mother of Akhenaten and grandmother of Tutankhamun, and the KV55 mummy, who is most probably Akhenaten, father of Tutankhamun (Figure 2, eAppendix, and online interactive kinship analysis and pedigree here). The latter kinship is supported in that several unique anthropological features are shared by the 2 mummies and that the blood group of both individuals is identical.31-32



Disease or Amarna Artistic Style?

Macroscopic and radiological inspection of the mummies did not show specific signs of gynecomastia, craniosynostoses, Antley-Bixler syndrome or deficiency in cytochrome P450 oxidoreductase, Marfan syndrome, or related disorders (eAppendix, Table 2). Therefore, the particular artistic presentation of persons in the Amarna period is confirmed as a royally decreed style most probably related to the religious reforms of Akhenaten. It is unlikely that either Tutankhamun or Akhenaten actually displayed a significantly bizarre or feminine physique.

It is important to note that ancient Egyptian kings typically had themselves and their families represented in an idealized fashion. A recent radiographic examination of the Nefertiti bust in the Berlin Museum illustrates this clearly by showing that the original face of Nefertiti, present as a thin layer beneath the outer surface, is less beautiful than that represented by the artifact.33 Differences include the angles of the eyelids, creases around the corners of the mouth on the limestone surface, and a slight bump on the ridge of the nose.34 Thus, especially in the absence of morphological justification, Akhenaten's choice of a "grotesque" style becomes even more significant.



Walking Impairment and Canes

Tutankhamun had a juvenile aseptic bone necrosis of the left second and third metatarsals (Köhler disease II, Freiberg-Köhler syndrome). The widening of the metatarsal-phalangeal joint space, as well as secondary changes of the second and third metatarsal heads, indicate that the disease was still flourishing at the time of death.35 Bone and soft tissue loss at the second metatarsal phalangeal articulation could further indicate that an acute inflammatory condition was present on the basis of an ulcerative osteoarthritis and osteomyelitis. The congenital equinovarus deformity (pes equinovarus) together with the malformed second toe of the left foot (oligodactyly [hypophalangism]) transferred additional joint load to the right foot, causing flattening of the foot arch (pes planus).

There is evidence that Tutankhamun may have had this impairment for quite some time. The walking disability can be substantially aided by the use of a cane. Howard Carter discovered 130 whole and partial examples of sticks and staves (eFigure 3A) in the king's tomb, supporting the hypothesis of a walking impairment.36 Traces of wear can be seen on a number of the sticks, demonstrating that they were used in the king's lifetime (eFigure 3B). Additional evidence for some sort of physical disability is found in a number of 2-dimensional images from Tutankhamun's reign that show him seated while engaged in activities for which he normally should have been standing, such as hunting (eAppendix and eFigure 3C).37-38



Malaria Tropica

Macroscopic studies revealed areas of patchy skin changes on the pharaoh's left cheek and neck of uncertain anamnesis, possibly indicating an Aleppo boil, a plague spot, an inflamed mosquito bite, or a mummification artifact.39 However, the genetic identification and typing of plasmodial DNA in Tutankhamun, Thuya, Yuya, and TT320-CCG61065 showed that they must have had malaria tropica, the most severe form of malaria (eAppendix).

Literary evidence for malaria infection dates back to the early Greek period, when Hippocrates described the periodic fever typical of this disease.40 Although it is believed that malaria widely affected early populations before Hippocrates,27, 41 until now only 1 report using immunological tools42 and few molecular genetic studies have clearly identified P falciparum in ancient specimens.43-46 We not only identified this parasite in our sample but also observed individual differences in some of the gene sequences as well as different MSP1 allele constellations in the 4 positive mummies. The diversity of plasmodial DNA (ie, variability in the genes' base order, length polymorphisms, or both) is a well-known phenomenon; however, some of the base deviations were not found in current DNA databases. Further research is required to typify these alterations in more detail and to assign these potentially unknown patterns to ancient Egyptian Plasmodium strains that date back to 3300 to 3400 years before present.

To our knowledge, this is the oldest genetic proof for malaria in precisely dated mummies. When the infection occurred, its severity, and whether it could have caused the death in the 4 mummies testing positive is not known. Preliminary data show that Tutankhamun and Yuya had multiple infections, as could be seen by the presence of the 2 P falciparum alleles MAD20 and RO33 of the MSP1 in the extracts. In contrast, and taking only the MSP1 test system into account, Thuya was infected by only 1 strain, which displayed the RO33 allele.

To date, no association has been found between P falciparum MSP1 genotypes and the clinical status of persons affected.47 We note that mixed P falciparum infections were detected in up to 78% of a contemporary sampling, and even isolates from symptomatic children contained more than 1 Plasmodium clone.47-48 Thus, multiple infections appear to be the norm rather than the exception. Moreover, the MSP1 allele frequencies tend to vary largely in different, sometimes even neighboring, areas but also over time.29 Thus, the prevalence rate of infection is not known—nor is it known if malaria was an epidemic or an endemic disease and how widely it was distributed in ancient Egypt.

Unfortunately, there is also no distinct evidence in ancient Egyptian texts of treatments for malaria, and there are no references to the fevers and chills associated with the disease.49 However, the Nile Delta and the fringes of the Nile Valley were marshy areas and thus excellent breeding grounds for the mosquito genus Anopheles. Interestingly, mosquitoes are mentioned in at least 1 ancient text,50 and it has also been suggested that the wooden frame of Queen Hetepheres (fourth dynasty) served as the support for a mosquito net.50 Herodotus also mentions that Lower Egypt was infested with mosquitoes or other insects and that people slept under nets to avoid them.51 Since there is nothing in the historical or archeological record that speaks against the widespread presence of this carrier in Pharaonic times, there is no evidence that can be used to argue against the diagnosis of malaria.



Cause of Death

Caution must be taken when interpreting cause of death in these mummies. It can be speculated that Yuya and Thuya had malaria, but it is not known if this was lethal (Table 3). Surprisingly, both individuals had reached an advanced (for the time) age of approximately 50 years or older (Table 1). This means either that the infection took place quite late in their lifetime, that they enjoyed strong genetic fitness, or that they aquired a partial immunity against the pathogen during their lives. Not every person infected with P falciparum becomes gravely ill, and this is especially true in populations that have been exposed to malaria pathogens over long periods.52 If Yuya and Thuya spent much of their time living in malaria-endemic areas close to the marshes of the Nile River, partial immunization may have contributed to their survival.

On the other hand, Tutankhamun had multiple disorders, and some of them might have reached the cumulative character of an inflammatory, immune-suppressive—and thus weakening—syndrome (Table 3). He might be envisioned as a young but frail king who needed canes to walk because of the bone-necrotic and sometimes painful Köhler disease II, plus oligodactyly (hypophalangism) in the right foot and clubfoot on the left. A sudden leg fracture23 possibly introduced by a fall might have resulted in a life-threatening condition when a malaria infection occurred. Seeds, fruits, and leaves found in the tomb, and possibly used as medical treatment, support this diagnosis (eAppendix, eFigures 3D and 3E).24-25,53-57

In conclusion, this study suggests a new approach to research into the molecular genealogy and pathogen paleogenomics of the Pharaonic era. With additional data, a scientific discipline called molecular Egyptology might be established and consolidated, thereby merging natural sciences, life sciences, cultural sciences, humanities, medicine, and other fields.
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PostPosted: Wed Feb 17, 2010 2:27 am    Post subject: Reply with quote

AUTHOR INFORMATION

Corresponding Author: Carsten M. Pusch, PhD, Institute of Human Genetics, Division of Molecular Genetics, University of Tübingen, Wilhelmstraße 27, D-72074, Tübingen, Germany (carsten.pusch@uni-tuebingen.de<!--
var u = "carsten.pusch", d = "uni-tuebingen.de"; document.getElementById("em0").innerHTML = '<a href="mailto:' + u + '@' + d + '">' + u + '@' + d + '<\/a>'//-->).

Author Contributions: Drs Hawass, Gad, Zink, and Pusch had full access to all of the data in the study and take responsibility for the integrity of the data and the accuracy of the data analysis.

Study concept and design: Hawass, Gad, Zink, Pusch.

Acquisition of data: Hawass, Gad, Ismail, Khairat, Fathalla, Hasan, Ahmed, Elleithy, Gaballah, Wasef, Fateen, Amer, Gostner, Selim, Zink.

Analysis and interpretation of data: Hawass, Gad, Ismail, Khairat, Fathalla, Hasan, Ball, Wasef, Fateen, Amer, Gostner, Selim, Zink, Pusch.

Drafting of the manuscript: Hawass, Gad, Zink, Pusch.

Critical revision of the manuscript for important intellectual content: Hawass, Gad, Ismail, Khairat, Fathalla, Hasan, Ahmed, Elleithy, Ball, Gaballah, Wasef, Fateen, Amer, Gostner, Selim, Zink, Pusch.

Statistical analysis: Ball, Gostner, Zink, Pusch.

Administrative, technical, or material support: Hawass, Gad, Ismail, Hasan, Ahmed, Elleithy, Ball, Gaballah, Fateen, Amer, Selim, Zink, Pusch.

Study supervision: Gad, Ismail, Zink, Pusch.

Financial Disclosures: None reported.

Funding/Support: This study was supported by the Discovery Channel and the Brando Quilici production group. Funding was also obtained from the Mini-Graduiertenkolleg Tübingen and the DAAD (GERLS exchange program). Siemens medical donated material and installed the multislice computed tomography scanner used in the study.

Role of the Sponsor: The funding organizations had no role in the design and conduct of the study; the collection, analysis, and interpretation of the data; or the preparation, review, or approval of the manuscript.

Online-Only Material: The eAppendix, eFigures 1 through 3, and the online feature showing full-body computed tomography reconstructions of the mummies and interactive kinship analysis and pedigree are available here.

Additional Contributions: Jessica Sherry, BS (manager, first laboratory), and Milena Gozzo, BDM (manager, second laboratory), contributed expert management of the ancient DNA laboratories in Cairo, Egypt. Neither of these individuals received compensation for their contributions beyond their salaries.

This article was corrected online for typographical errors on 2/16/2010.

Author Affiliations: Supreme Council of Antiquities, Cairo, Egypt (Dr Hawass and Mr Elleithy); National Research Center, Cairo, Egypt (Drs Gad, Ismail, and Amer and Mss Hasan and Ahmed); Ancient DNA Laboratory, Egyptian Museum, Cairo, Egypt (Drs Gad and Ismail and Mss Fathalla, Khairat, Hasan, and Ahmed); Institute of Human Genetics, Division of Molecular Genetics, University of Tübingen, Tübingen, Germany (Ms Khairat, Mr Ball, and Dr Pusch); Learning Resource Center, Kasr Al Ainy Faculty of medicine, Cairo University, Cairo, Egypt (Drs Gaballah and Fateen and Ms Wasef); Department of Radiodiagnostics, Central Hospital Bolzano, Bolzano, Italy (Dr Gostner); Department of Radiology, Kasr Al Ainy Faculty of medicine, Cairo, Egypt (Dr Selim); and Institute for Mummies and the Iceman, EURAC, Bolzano, Italy (Dr Zink).
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PostPosted: Wed Feb 17, 2010 2:29 am    Post subject: Reply with quote

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30. Mlambo G, Sullivan D, Mutambu SL; et al. Analysis of genetic polymorphism in select vaccine candidate antigens and microsatellite loci in Plasmodium falciparum from endemic areas at varying altitudes. Acta Trop. 2007;102(3):201-205. FULL TEXT | WEB OF SCIENCE | PUBMED

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35. Diethelm L, Olsson O, Strnad S, Vieten H, Zuppinger A. Handbuch der medizinischen Radiologie V/4, Skeletterkrankungen. Berlin, Germany: Springer; 1976.

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41. Joy DA, Feng X, Mu J; et al. Early origin and recent expansion of Plasmodium falciparum. Science. 2003;300(5617):318-321. FULL TEXT | WEB OF SCIENCE | PUBMED

42. Bianucci R, Mattutino G, Lallo R; et al. Immunological evidence of Plasmodium falciparum infection in an Egyptian child mummy from the Early Dynastic Period. J Arch Sci. 2008;35(7):1880-1885.

43. Sallares R, Gomzi S. Biomolecular archaeology of malaria. Anc Biomol. 2001;3:195-213.

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PostPosted: Wed Feb 17, 2010 2:38 am    Post subject: Reply with quote

---END---

//---there is additionally an eSuppliment which came in .pdf format (much the same information but with some additions (pics &c.). I dunno how to share this (unless someone can tell me) alternatively send me an email and I will forward the .pdf file, or I can forward it to one of the mods for distribution (whatever)---//

Anyway, most all of this will probably become ubiquitous online over the next short while, but I wanted to share with folks who have a real interest

Now perhaps the board can enjoy some real interesting discussion Idea
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PostPosted: Wed Feb 17, 2010 3:07 am    Post subject: Reply with quote

Thanks for sharing that freeThinker.
It will take a while until I can actually absorb all the info Very Happy

Anneke
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PostPosted: Wed Feb 17, 2010 3:20 am    Post subject: Reply with quote

This stuff is slightly interesting news, eh?

Understatement of the year. Very Happy

I got the EEF posting and have read through some of the links there and some of the posts here, and I have a couple of questions. I apologize if the answers to my questions have already been covered here, but I'm short on time and will check in again later.

First, the KV55 mummy has been identified as the father of Tutankhamun. How do we know that this is Akhenaten, however? It was my understanding that the skeleton in KV55 was most likely that of a male no more than 25 years of age at death, so too young to have been Akhenaten.

Second, the mummy in KV21 might be Ankhesenamun. I don't know much of anything about this mummy, but if memory serves, two female mummies were found in that tomb and both have been thought to be possible royal ladies in Dynasty 18. And that right there is probably the sum total of my knowledge about the mummy in question.

So my question about her is, has it been established in these genetic analyses that the KV21 mummy was the daughter of the KV55 mummy? Is that why KV55 is being identified as Akhenaten? I understand that it is now established that KV21 was the mother of the two stillborn girls in KV62, so I'd love to know if KV21 can be genetically tied to KV55.

And to echo anneke in one of her first posts, who on earth might KV35YL be? That's especially intriguing to me, as the mother of Tutankhamun (I mean the mummy--I'm not Tut's mom). I don't think there's much chance that either Kiya or Nefertiti were a full sister to Akhenaten, is there?

Thanks to anyone and everyone who can shed light on these questions. I look forward to learning more. Smile
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PostPosted: Wed Feb 17, 2010 3:45 am    Post subject: Reply with quote

It seems, the 'film of the book' is to be shown on Sunday. Discovery 8pm EST (sorry dunno what is in store for folks across the pond). Perhaps we gotta watch that before getting rewarded with ACT III

I had same question about KV55=Akhenaten - heck! we're back at Arthur Weigall - seems like almost full circle Razz
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PostPosted: Wed Feb 17, 2010 4:06 am    Post subject: Reply with quote

I am wondering the same thing about the identity of KV 55.

It seems KV 55 and KV 35YL are a bother and sister and the parents of Tut, and also the children of Amnhotep III and Tiye. Not sure what the possibilities are besides maybe Akhenaten and a sister or maybe Smenkhare and a sister (Nebetah, Baketaten?).

As the father of Tutankhamen and the son of Amenhotep III and Tiye, it seems that this could be Akhenaten. But if that is the case then either the individual is at the higher end of the age estimate - maybe we can stretch it to late 20s? This would have some implications re the age of Akhenaten when he came to the throne: ie most likely a minor? Is this why some of the foreign rulers are corresponding with his mother?

If KV 55 is Akhenaten, then he married a full sister which begs the question why a daughter of Queen Tiye - as powerful as she was - did not play a more prominent role. Unless this means though that Nefertiti was a daughter of Amenhotep and Tiye. A long time ago some did interpret one of scenes showing Tiye's visit as actually saying she was the mother of both Akhenaten and Nefertiti. It was a bit of a controversial reading of the inscription and it is now not read that way. Maybe this should be re-examined?

Or this person could be Smenkhare? Then Smenkhare would be the son of Amenhotep III and Tiye and as such the third son (after Tuthmose and Akhenaten). And Smenkhare may have married one of the younger sisters? Maybe Nebetah or Baketaten? But Smenkhare only appears as the husband of Meritaten. So that is a bit odd. We know for sure she was the daughter of Nefertiti. Unless this brother-sister marriage dates to an earlier time? Would this be a reason for the death of the mother of Tut?

If KV 55 is Smenkhare, then Akhenaten must have died without an heir and the throne went to his brother. Which raises more questions about who Neferneferuaten was. Was this Nefertiti trying to hold on to power? Function as a co-regent with Akhenaten?

But if Tut really married his sister, then he must have been the son of Akhenaten. It seems to me that the problems with the sequencing of the DNA of KV 21 A is a bit of an issue. She seems to be the mother of the stillborn children, but her lineage is not certain.
Could Tutankhamen have married his cousin?

LOL It seems that this information raises as many questions as it answers.

It seems that no matter what we have some interesting power transfers.


All I can say is that it really became even more confusing ....
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PostPosted: Wed Feb 17, 2010 4:40 am    Post subject: Reply with quote

This from the supplement .pdf that came with material posted above:

A probability of more than 99.7% is regarded as "paternity is practically proven"

RESULTS
Kinship Analyses

... The statistical analysis revealed that the mummy KV55 is most probably the father of Tutankhamun (probability of 99.99999981%) and KV35 Younger Lady could be identified as his mother (99.99999997%). The testing of Amenhotep III as father of Tutankhamun and KV35 Elder Lady as putative mother were both negative owing to mismatching alleles. Amenhotep III could clearly be identified as father of KV55, showing a paternal probability of 99.99999999%. The results demonstrate that the mummy in KV55 is the son of Amenhotep III and father of Tutankhamun, leading to the assumption (also supported by the radiological findings) that the mummy can be identified as Akhenaten. It could be further shown that Tutankhamun is the most likely father of the 2 fetuses found in KV62 (Fetus1: 99.97992885%, Fetus2: 99.99999299%). The degree of shared alleles between the female mummy KV21A and Fetus1 and Fetus2 points toward a possible identification of the mummy as Ankhensenamun, the mother of both fetuses and wife of Tutankhamun...
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kylejustin
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PostPosted: Wed Feb 17, 2010 4:41 am    Post subject: Reply with quote

from what i understand, tuts babies are mitochondrial descendants of thuya. so if ankhsenamun is their mother, then nefertiti is a female relation of queen tiye. but, if the kv 21 mummy is their mother, why doesnt she have the same mitochondrial dna? anksenamun could still be the mother, but this mummy doesnt have the same mitochondrial dna as the babies from what i understand? i got this from the dna chart on the first page of this thread.

i also think akhenaten isnt his father. both mummies identified as tuts parents are supposed to have died in their early 20's. which is really throwing out aidan dodson's theories. so we can eliminate akhenaten's daughters as tuts mother. so we are left with a full blood sister of akhenaten. i think smenkhare is still front runner for kv 55. therfore he will be tuts father. he ruled after akhenaten which makes sense if tut was a) not akhenaten's son, or b) if he was too young to rule, and smenkhare was supposed to be regent.

tuts mum has very violent injuries, as stated in the above table about the mummies. which one of akhenatens sisters could be that much of a threat?

also, are we to think that the mummy who had an arrow wound in his chest is the one we usually identify as thutmose I?

and are we to understand that kyphoscoliosis is a gentic trait in the 18th dynasty? since thutmose II and sitre in had it? are we to now think sitre in, the nurse of hatshepsut, is now a member of the thutmosid bloodline?
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christphe
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PostPosted: Wed Feb 17, 2010 6:02 am    Post subject: Reply with quote

[quote="anneke"]I am wondering the same thing about the identity of KV 55.

As the father of Tutankhamen and the son of Amenhotep III and Tiye, it seems that this could be Akhenaten. But if that is the case then either the individual is at the higher end of the age estimate - maybe we can stretch it to late 20s? This would have some implications re the age of Akhenaten when he came to the throne: ie most likely a minor? Is this why some of the foreign rulers are corresponding with his mother?

If KV 55 is Akhenaten, then he married a full sister which begs the question why a daughter of Queen Tiye - as powerful as she was - did not play a more prominent role. Unless this means though that Nefertiti was a daughter of Amenhotep and Tiye. A long time ago some did interpret one of scenes showing Tiye's visit as actually saying she was the mother of both Akhenaten and Nefertiti. It was a bit of a controversial reading of the inscription and it is now not read that way. Maybe this should be re-examined?

quote]

in is book Marc Gabolde raise some "evidences" that amenhotep IV was very young when he became king. This is why he is seen with is mother in Keruef tomb and this explain some letters found in amarna like if tiyi was a "regent"or at least an advisor. He also raise the idea that his majority is celebrated by marrying Nefertiti. So in fact he was young when he died.
I have never read that Nefertiti was his sister, this would explain her exeptional religious position.

If Ankhsenamon body is identified than it should be easy to now if tut mother is Nefertiti or another woman. What about Baketaton?
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kylejustin
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PostPosted: Wed Feb 17, 2010 6:51 am    Post subject: Reply with quote

this can probably be discount right away, but is it possible the kv 55 mummy is crown prince thutmose? i just find it very hard to believe the mummy is akhenaten's. and if it is, why was smenkhare in the co regency before tutankhamun? unless smenhkare is also akhenaten's son?

i just think that if the bones havnt fused and the teeth havnt developed, how can he be more than 20-25 at death?
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PostPosted: Wed Feb 17, 2010 7:05 am    Post subject: Reply with quote

kylejustin wrote:
...i just find it very hard to believe the mummy is akhenaten's. and if it is, why was smenkhare in the co regency before tutankhamun? unless smenhkare is also akhenaten's son?


Perhaps; just as KV55 had been >>> proposed and then >>> rejected and now >>> accepted as Akhenaten - so might Smenhkare having been previously >>> proposed and then rejected >>> and now (?) back to being just a name change (alternative) for Nefertiti (? - as co-regent?)

We have no body for smenhkare now, so would it not be fair to say... we have no real gender?

Maybe the ephemeral smenhkare has just dissapeared in a puff of smoke. I'll bet all this ends up with discussion about smenhkare rather than tut or akhenaten- hmmm...
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christphe
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PostPosted: Wed Feb 17, 2010 7:20 am    Post subject: Reply with quote

during XVIII dynasty kingship didn't pass from brother to brother but father to son. That"s probably why we hear about princess but not princes. this may explain why Ramses II introduced changes by promoting his sons.
The idea of Smenkhare bother of akhenaton brother of tuthankamon would appear out of tradition.
It seems that all was made to secure Tut accession to the throne in spite of his age, the same way they did with thoutmose III or Amenhotep III.
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