After 3,400 years, one of history’s oldest cold cases has finally been solved.

In addition to finding evidence that the legendary Egyptian pharaoh Tutankhamun (better known as King Tut) died of malaria and suffered from a club foot, DNA analysis unveiled last week has identified a previously anonymous excavated mummy, KV55, as Tutankhamun’s father: Akhenaten, the rebellious pharaoh of Egypt’s Amarna period whose legacy disappeared for thousands of years. The DNA supports the theory of two Yale professors, who suggested that the androgynous figures of the period’s art reflect religious beliefs rather than representations of Akhenaten’s family.

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This theory was presented by Yale Egyptology professors John Darnell and Colleen Manassa ’01 GRD ’05 in their 2007 book, “Tutankhamun’s Armies,” which details the military and diplomatic technique of Egypt’s late 18th Dynasty, which ruled the country from 1550 B.C. to 1292 B.C.

KV55’s name refers to the location of Akhenaten’a tomb in the Valley of the Kings, a burial site near Thebes, or modern-day Luxor. The tomb was excavated by archaeologist Edward Ayrton in 1907, and until last week, the identity of its inhabitant had remained a mystery, Darnell said. The study to discover the identity of KV55 was lead by archaeologist Zahi Hawass, the current secretary general of the Egyptian Supreme Board of Antiquities, and was chronicled on a Discovery Channel program, “King Tut Unwrapped,” which aired earlier this week. Darnell and Manassa had been invited by Hawass to discuss military aspects of Akhenaten’s and Tutankhamun’s rule on the show.

“It was a neat way to learn more about ancient Egypt in a way just we didn’t have access to before,” Manassa said.

Previously, genetic material had not been available to confirm Tutankhamun’s cause of death — which was once believed to be a blow to the head — or the theories in “Tutankhamun’s Armies,” which were based on archeological excavations and textual examinations, Manassa said. Because the DNA results ruled out that Akhenaten had a congenital condition such as Marfan’s syndrome, a genetic disorder that leads to a tall and thin figure and slender limbs, they vindicated Darnell and Manassa’s belief that the androgynous figures depicted in the art of the period were shown that way for religious reasons rather than to represent Akhenaten’s physical deformities.

Akhenaten’s rule, now know as the Amarna period in reference to the capital city’s movement to Amarna, was a period of upheaval in Egyptian history, Manassa said. Akhenaten and his queen had ruled as though they were the son and daughter of the Sun God Aten, and the androgynous depictions of Amarna art — rotund stomachs, feminine hips and breasts — were meant to call these divine beings to mind, Darnell said.

Manassa said the Amarna period has been misinterpreted on many counts.

“It was an obscure period with great popularity, which gave rise to many incorrect theories,” Darnell said. “If these results move people towards focusing on fact, that will be a great thing.”

Darnell and Manassa are currently on sabbatical working on another project. In the spring of 2011, they will teach a class titled “Age of the Amarna” with Karen Foster, a lecturer on the art of the ancient Near East and Aegean.