You remember these famous words from Shakespeare:
Now is the winter of our discontent
Made glorious summer by this sun of York;
And all the clouds that lour’d upon our house
In the deep bosom of the ocean buried.
They are of course the opening lines of “Richard III,” and are spoken by the nefarious and hunchbacked king. Although he’s called “Gloucester” in the play’s text, he was a real person—King Richard III—and he was deformed, having scoiliosis (Richard’s deformity is confirmed by historical records, including descriptions by Thomas More).
I could simply rehash the history of this short-lived king (1452-1485), but I’ll let the authors of a new paper in Nature Communications (reference and link at bottom; free download) give you the backstory. For these are the scientists who identified his body, found underneath a carpark in Leicester two years ago; and they published what seems to be a definitive ID based on skeletal, dating, and DNA-based evidence. The genetic evidence is way cool, for besides identifying him, they could also get a good idea of what his hair and eye colors were, and check those against the historical record. King et al. say this about Richard:
Richard III is one of the most famous and controversial English kings. His ascension to the throne in 1483, following the death of his brother, Edward IV, has been seen as contentious, involving, as it did, discrediting the legitimacy of Edward’s marriage and therefore the claim of both of Edward’s sons to the throne. Later, as yet unproven accusations arose that Richard had his two nephews murdered to solidify his own claim. Richard’s death two years later on August 22nd 1485 at the Battle of Bosworth marked the end of the Plantagenet dynasty, which had ruled for over 300 years, and the beginning of the Tudor period. Richard III was the last English king to be killed in battle, he became one of Shakespeare’s most notorious villains, and is one of the few English monarchs whose precise resting place was lost: the mystery surrounding the fate of his remains persisting to the present day.
Historical records report that after Richard III was killed on the battlefield, age 32, his remains were brought back to Leicester and buried in the medieval church of the Grey Friars. The friary was dissolved in 1538 under the orders of King Henry VIII, with most of the buildings being torn down in the following years. Approximately 125 years later, a rumour arose that Richard III’s remains had been disinterred during the dissolution of the monasteries and thrown into the river Soar in Leicester. However, it had long been thought that this rumour was unsubstantiated and it was therefore expected that the grave of Richard III should still lie within any remains of the Grey Friars church. While historical records and the subsequent analysis thereof have long indicated the approximate location of the Grey Friars friary, and its likely situation in relation to the modern urban landscape of Leicester, the exact site of Richard III’s grave had been lost in the 527 years since his death.
Although Richard III reigned for only a little over two years, substantial historical information about various features of his life and death exists. These include aspects of his physical appearance such as having a slim build, one shoulder higher than the other and that he suffered battle injuries, which resulted in his death. In September 2012, a skeleton (Skeleton 1) was excavated at the presumed site of the Grey Friars friary in Leicester, the last-known resting place of Richard III.
How can you ID bones from remains that are over four centuries old? Well, first you see if the skeleton looks like someone who could be Richard III. And indeed it did:
Male skeleton, aged 30-34. Check.
Slim build, scoliosis (one shoulder higher than the other). Check.
Carbon dating consistent with historical data. Check: dated from 1456-1530 with 95.4% probability.
Skeletal injuries consistent with death in battle. Check.
That’s pretty good, for there can’t be many swaybacked people who died in battle at the right time, and of the right age, buried where Richard is rumored to have been buried. But to clinch the case, the authors did DNA analysis, extracting the genetic material from the skeleton’s teeth and bones. They used three types of DNA, sequencing them and then seeing if they matched with the known descendants of Richard (yes, there are some, for British royal genealogies were kept scrupulously).
Here, if you’re interested, are the genealogies of the several relatives of Richard whose DNA was matched to that of his putative remains (click to enlarge):
They used three types of DNA, and I’ll give the results separately.
1. Mitochondrial DNA (mtDNA). This is the DNA in the mitochondrion, a cellular organelle that produces energy and has its own DNA molecule distinct from the “regular” DNA in the nucleus. Since it’s in the cell’s “cytoplasm” (the contents of the cell), mtDNA is passed on solely from mother to offspring, as all the cytoplasm in a fertilized egg comes from the mother. The father’s mitochondrial DNA is not inserted into the egg that is fertilized by his sperm, so that part of his DNA isn’t present in his kids.
Thus, if you’re matching mtDNA of the putative Richard III with his descendants, you need to find an unbroken lineage of female descendants that come from either Richard’s mother or his sister. And, fortunately, that existed. If you look at the right-hand side of the figure above, you can see that Richard’s oldest sister, Anne of York, has an unbroken lineage of female descendants giving rise to two living people, Michael Ibsen (a male) and Wendy Duldig (a female), who are now 14th cousins of each other. Since Anne of York and Richard III had the same mother, they would also have had the same mitochondrial DNA, and that would have been passed down to both Ibsen and Duldig.
And indeed, the mtDNA of these two living people matched nearly perfectly with that of the mtDNA from the skeleton. They sequenced the entire mtDNA (once a very hard task), and one of the relatives matched absolutely perfectly, base for base among about 16,500 bases. The other modern relative differed in only a single base—very likely a mutation that occurred in the intervening four centuries. The probability of finding such a perfect match between the skeletal sequence and a random modern British person (calculated from a British mtDNA database) was conservatively calculated at 2 in 1832. Of course this probability decreases if you allow for that one mutational difference in the other descendant. For all practical purposes, this is very strong evidence that Richard III was an ancestor of the two living descendants shown on the right above.
2. Y chromosome DNA. Y chromosomes are passed only from father to son, so to match the putative Richard III’s Y DNA with that from living people, you need an unbroken string of male descendants of Richard. We have that, too, and it’s shown on the left side of the figure above. In fact, they found five descendants (not named). What they found here, though was that they did not match Richard’s skeletal DNA!! Now you could say that this means that Richard was not their ancestor at all, but another hypothesis (more likely given the forensic skeletal and mtDNA matches) is that there was what evolutionist John Maynard Smith called some “sneaky fucking” in the British royal family that mixed some non-royal DNA with the royal DNA.
That is, somewhere fairly far back in the lineage (for four of the living male relatives had the same DNA), some male who was supposedly a descendant of Richard III was actually fathered by someone outside the lineage. In other words, a British royal female had a bit of a fling. Those liaisons are harder to detect when you use mtDNA (which makes the mtDNA more reliable for forensics like those in this survey), for while you always know who the mother of a child is (after all, she gave birth to it), you aren’t always sure about the father. (This is called, of course, “paternity uncertainty”.) But Y chromosomal DNA is good for determining in modern cases whether a male really did father a child.
The authors note that paternity uncertainty for a given child is about 1-2%, and calculate that in the many generations between Richard III and modern descendants, the probability that some sneaky fucking occurred in that lineage would be around 16%. It could of course be higher or lower depending on the frequency of royal flings.
The implications of this, given the genealogy of Brits, are potentially large, for they could mean that a big swath of historical British royalty was genealogically bogus! As the authors note (read this carefully, my emphasis):
One can speculate that a false-paternity event (or events) at some point(s) in this genealogy could be of key historical significance, particularly if it occurred in the five generations between John of Gaunt (1340–1399) and Richard III (see Supplementary Fig. 2). A false-paternity between Edward III (1312–1377) and John would mean that John’s son, Henry IV (1367–1413), and Henry’s direct descendants (Henry V and Henry VI) would have had no legitimate claim to the crown. This would also hold true, indirectly, for the entire Tudor dynasty (Henry VII, Henry VIII, Edward VI, Mary I and Elizabeth I) since their claim to the crown also rested, in part, on their descent from John of Gaunt. The claim of the Tudor dynasty would also be brought into question if the false paternity occurred between John of Gaunt and his son, John Beaufort, Earl of Somerset. If the false paternity occurred in either of the three generations between Edward III and Richard, Duke of York, the father of Edward IV and Richard III, then neither of their claims to the crown would have been legitimate.
But since we don’t know when or where the surreptitious insemination occurred, we can’t say for sure that the House of Tudor was indeed genealogically bogus.
And, of course, this doesn’t affect the genealogy of the present British family tree, which are Windsors, for the Tudor family tree died out and was replaced by the Stuarts, Hanovers, and then the present Windsors (formerly the House of Saxe-Coburg and Gotha; the name was changed in 1917 to prevent the Brits from sounding German). So, to my chagrin (I don’t like British royalty), Elizabeth and her offspring and her offspring’s offspring can still claim royalty.
3. Autosomal DNA. Most of our DNA is carried on the autosomes, or the set of 44 chromosomes (22 pairs) that doesn’t determine sex (that’s the other pair, either XX or XY). And we know from sequencing the human genome and from “association mapping” which regions of the autosomal genome play major roles in determining hair and eye color. If you sequence those parts of the autosomal DNA, then, you can get a good idea about a person’s hair and eye color even without seeing it.
And they did this for those parts of the autosomal DNA of the putative Richard III skeleton. Although there are no contemporary portraits of Richard III, one painting in the 1510s (the “SAL painting”) was made about 25 years after his death, and it’s shown below. It shows the king with blue eyes and brownish hair.
DNA typing from the autosomes shows that Richard had blue eyes with a probability of 96% and blond hair with a probability of 77%. However, those same genotypes can produce a variety of hair colors, including blond darkening to brown with age, as shown in the top part of the figure below. (All of these living individuals have the same forms of genes for hair and eye color found in the Richard III skeleton). Of all the existing paintings of Richard III, the one that most closely matches these traits is also the earliest one and the one deemed most authentic, the SAL painting.
The upshot: The authors combined all the probabilities: genetic (including the mismatch of the Y chromosomes but also the possibility of false paternity), skeletal, and dating, to calculate a likelihood that the skeleton really was that of Richard III. To do this they had to incorporate a priori estimates that the skeleton was his, and used two: a “skeptical” probability (2.5%) and a more liberal one (50%). Combining all these, they get a probability between 99.9994% and 99.99999% that the skeleton found was in fact that of Richard III. These are conservative probabilities, so for our purposes we can in fact be dead certain that the skeleton found was that of Richard III.
Envoi
Finally, according to a BBC story from yesterday, Richard’s remains will be reburied in the Leicester Cathedral on March 26 of next year, in this tomb:
They also provide a photo of Richard’s skull next to another painting of him, a reminder of our mortality (as if we needed it!):
UPDATE: A reader below points to a University of Leicester post showing a model of what Richard III would have looked like, based on the skull. They get the hair and eye colors wrong, for that information wasn’t known when the model was made. And it doesn’t look a lot like the painting. But what I really like on the site is a reconstruction of what Richard’s speech might have sounded like, based on his writing and on reconstruction of pronounced English of that time. It is very different from modern English!
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King, T. E. et al. 2014. Identification of the remains of King Richard III. Nature Communications, published online Dec. 2; DOI: 10.1038/ncomms6631
