STATE-of-the-art genetic testing has now proved beyond question that the remains found under a council car-park two years ago were those of England’s last Plantagenet king.

Analysis of all the evidence has confirmed the bones as those of Richard III to the point of 99.999 per cent – “at its most conservative.”

It also shows the king was almost certainly blue-eyed and probably had blond hair, at least during his childhood.

And, importantly for historians, it has also provided scientific evidence for the first time of a question mark over medieval lines of succession in the monarchy.

It shows the male line of descent was broken at one or more points between Richard and his living relatives.

Historically Royal succession has taken a number of twists and turns over the centuries and is not based on straight linear inheritance.

But depending on where the break occurred it could have potential implications for the historical monarchy – although the likelihood is it was well down the chain and therefore would not have affected Royal lineage.

And Professor Kevin Schürer of Leicester University insisted the results showed there was “absolutely no evidence that a link in the Royal chain of succession had actually been broken.”

The battle-scarred remains of Richard, who spent much of his childhood at Middleman also had close connections with York, were discovered in Leicester in August 2012 by experts from the city’s university.

The complex genetics and painstaking genealogical research that followed has now been published in full in the journal Nature Communications.

The paper is the first to carry out a statistical analysis of all the evidence together to prove beyond any reasonable doubt that skeleton from the site in Leicester is indeed the remains of Richard III.

The research team was led by Dr Turi King who said: “Even with our highly conservative analysis, the evidence is overwhelming that these are indeed the remains of King Richard III, thereby closing an over 500-year-old missing person’s case.”

Dr King was part-funded by the Wellcome Trust whose director culture and society Simon Chaplin said: “It is exciting to have access to genetic data from any known historical individual, let alone a king of England lost for more than 500 years.

“Adding this information to a wealth of existing material about Richard III further highlights the ways in which studying human remains can inform our understanding of the past, and we look forward to learning more about Richard for many years to come.”