14

Richard III’s Genes part I – the Fifteenth Century and Before

Modern attempts to discern ‘the real Richard III’ behind the ‘Tudor’ and later image have certainly encompassed an astonishingly wide variety of approaches and sources. These have included the study of fifteenth-century and later written evidence, examination of Richard’s portraits, and analysis of the books which comprised Richard’s library. In addition, attempts have been made to analyse Richard III’s handwriting, and to cast his horoscope.¹ However, prior to my research, no-one had attempted to explore Richard’s genetic make-up.

Before 2012, no direct investigation of Richard III’s physical remains was possible because, until the recent excavations in Leicester, Richard’s body had been inaccessible. Therefore I initially sought alternative sources of appropriate genetic material. The obvious route would have been the remains of one of Richard’s descendants, but this approach also appeared to be closed to us. The king’s only legitimate son, Edward, Prince of Wales, died in childhood, some months before his father. What some believe to be the boy’s tomb is in the church at Sheriff Hutton, in Yorkshire, but the identity of this burial is controversial, nor is it easy to obtain permission for the exhumation of royal remains. Richard’s other known children, Catherine Plantagenet and John of Gloucester, were illegitimate. John was executed by Henry VII.² His tomb and that of his (half?) sister have not been located. Moreover, since (so far as is known) all Richard III’s children died childless, the king apparently has no direct living descendants.³ Thus, in seeking Richard III’s genes I was obliged to range further afield, exploring collateral lines of descent from Richard’s siblings. Of potential interest were Richard’s parents, the Duke and Duchess of York; his brothers, Edward IV, Edmund, Earl of Rutland and George, Duke of Clarence; and his sisters, Anne, Duchess of Exeter, Elizabeth, Duchess of Suffolk and Margaret, Duchess of Burgundy.

The burial sites of some of these individuals were known, and their remains could very easily have been examined – had their exhumation ever been authorised. The Duke and Duchess of York, together with Richard’s brother, Edmund, Earl of Rutland, lie entombed in secondary and tertiary burials at Fotheringhaye church. Earlier, their tombs were in the chancel, at the east end of the collegiate church, but when the Reformation led to the demise of the college and the ruin of the chancel, Elizabeth I had the bodies of her ancestors and relatives moved into the surviving nave of the building. The new tombs which she provided flank the present high altar.

Richard’s brother, King Edward IV, still lies in his tomb in St George’s Chapel, Windsor, and his eldest sister Anne of York, Duchess of Exeter, lies in a nearby side chapel, in the same building. Permission to exhume these remains is unlikely ever to be granted. Nevertheless, some genetic material from the body of Edward IV is available. This avenue has been explored, and we shall return to this point presently.

The burial site of Elizabeth of York, Duchess of Suffolk, the second surviving sister of Richard III, is also known. Elizabeth lies undisturbed in her alabaster tomb at Wingfield church in Suffolk. Apart from the king himself, she is one of the few members of Richard’s immediate family whose remains had lain untouched from the time of her death in 1503 until the present day.

As for George, Duke of Clarence, he was probably buried in Tewkesbury Abbey, Gloucestershire, but the remains once thought to be his are now considered to be of doubtful authenticity.⁴ These bones lie behind the high altar in what is known as the ‘Clarence Vault’. The remains are contained in a glass case which was opened and examined in 1982. On that occasion a physical examination revealed the fragmentary remains of at least two individuals – at least one male and at least one female. Superficially this sounds quite promising, since the Duchess of Clarence was unquestionably buried at Tewkesbury,⁵ and Clarence is believed to have been interred beside her. However, the male remains were tentatively assigned to the age range 40–60+ years, while the female remains were designated in the range 50–70+ years. This was completely incompatible with the known ages of the Duke and Duchess of Clarence at the time of their deaths (when they would have been 28 and 25 years of age respectively). It is known that in the eighteenth century the Clarence Vault was taken over for burials of members of the family of Samuel Hawling, an alderman of Tewkesbury, and it seems highly probable that at least some of the skeletal remains now in the Clarence Vault are those of Hawling and his wife and son, who were aged respectively 72, 96 and 86 at death.⁶ To date, however, no DNA testing of the bones from this vault has taken place. Osteological re-examination of the bones would be highly desirable and, depending on the results of that examination, DNA testing of them might at some stage be considered.

As for Richard’s youngest sister, Margaret of York, Duchess of Burgundy, the current whereabouts of her remains present major problems of their own.⁷ Nevertheless, it was the confusion regarding Margaret’s burial that provided the immediate impetus for my research which finally led to the establishment of a mitochondrial DNA sequence for Richard III and his brothers and sisters.

At her own request, Margaret of York’s body was buried in the Fransciscan Priory Church at Mechelen (Malines), in modern Belgium.⁸ This building lies just to the west of Mechelen’s cathedral church of St Rombout. The Priory Church, sacked during the religious conflicts of the ensuing centuries, is now a cultural centre, and all trace of Margaret’s once splendid tomb has vanished. A manuscript copy of Margaret’s memorial inscription tells us that she was buried ‘beneath the threshold of the doorway of this chancel’.⁹ This rather odd location, which now appears to reflect precisely the burial location of her brother Richard at the Franciscan Priory in Leicester, may not have been accidental. Perhaps Margaret deliberately requested burial in a priory church of the same order as her slaughtered brother, and deliberately asked for her tomb to be placed in an identical position, just inside the entry to the choir. Until Richard’s burial was discovered, however, the exact meaning of the somewhat imprecise description had been debated, owing to the fact that the choir of the Mechelen priory church may have had more than one entrance. Originally, of course, the meaning had presumably been clarified by the physical location of the bronze memorial plaque within the church. Sadly, however, this vital evidence was lost to us. As a result, doubts had been expressed as to where exactly Margaret’s corpse had been laid to rest.

In 2003 Dr Paul De Win published in Mechelen a paper on the multiple possible remains of Margaret of York.¹⁰ He explained the circumstances of Margaret’s burial, explored the subsequent vicissitudes of her tomb, and catalogued twentieth-century attempts to find her body. He also highlighted the problem of resolving which (if any) of the various female remains disinterred from the former Franciscan church in Mechelen might really be Margaret’s bones.

As reported in Paul De Win’s paper, three sets of female remains of approximately the right age were found in the former Franciscan church of Mechelen during the course of the twentieth century, and in locations which could potentially be interpreted as consistent with the approximate site of the lost tomb of Margaret of York.¹¹ These remains were found respectively in 1936 (excavations led by Vaast Steurs),¹² 1937 (excavations associated with the name of Maximilien Winders)¹³ and 1955 (accidental discovery, subsequently examined by Professor François Twiesselmann).¹⁴ Until recently these remains were stored in five boxes at the Mechelen Town Archives. They have recently been transferred to the Archaeology Service,¹⁵ and are now stored in two boxes (but reportedly a record of their former numbering has been retained). The bones from the 1955 discovery were photographed at the time, and these bones were subsequently coated with varnish. As a result, they can still be relatively easily identified. It is not currently possible to distinguish for certain which of the other female remains from Mechelen’s Franciscan Priory site were discovered in 1936, and which in 1937.¹⁶

In 2003, following discussions with Dr De Win, I began the attempt to establish a mitochondrial DNA sequence for Margaret of York and her siblings. Since mtDNA is normally inherited unchanged in the female line, the methodology adopted was to seek a living all-female-line descendant of Margaret’s mother, Cecily Neville, Duchess of York, or of one of Cecily’s close female relatives. We shall follow this research in a moment. First, however, it may be useful to summarise briefly what DNA is, and how it can currently be used in historical research.

The letters ‘DNA’ are an abbreviation for ‘deoxyribonucleic acid’. All living beings have DNA, which functions rather like an order pad. It lists, in coded form, the materials required to make the components of living bodies, and it specifies the order in which they must be assembled in order to create these components. In 1953 two Cambridge scientists, James D. Watson and Francis Crick, first worked out the structure of DNA, and demonstrated its significance as the basic coding material of life. ‘Watson and Crick had discovered that each molecule of DNA is made up of two very long coils, like two intertwined spiral staircases – a “double helix”. When the time comes for copies to be made, the two spiral staircases of the double helix disengage.’¹⁷

DNA has a very complicated molecular structure, but four principal components are the heterocyclic bases which are known by their initial letters: A for adenine, C for cytosine, G for guanine and T for thymine. In 1988, thirty-five years after the original discovery of DNA by Watson and Crick, an Oxford University team discovered that it was sometimes possible to extract, replicate and analyse DNA from ancient bones.¹⁸

While our present focus is on human DNA, the same basic rules apply to animals and plants, for all living things have DNA. Their cells contain two kinds of DNA: nuclear DNA, which resides in the cell nucleus, and mitochondrial DNA (mtDNA). Self-evidently, the latter is the DNA of the mitochondria: tiny structures which reside outside the cell nucleus in the surrounding cushion of cytoplasm and which help the cell to use oxygen in order to produce energy. The division in our cells between the two kinds of DNA is by no means equal. Each cell contains far more nuclear DNA than mitochondrial DNA. The latter represents a mere 0.5 per cent of our total.¹⁹

Nuclear DNA is a mixture, fifty per cent of which is inherited from each parent. Conversely, mitochondrial DNA is inherited from the mother, and is normally transmitted unchanged to the child. In addition, ‘mitochondrial DNA mutates at a much higher rate than nuclear DNA … Two organisms will therefore be far more similar in their nuclear DNA than in their mtDNA’.²⁰ For both of these reasons mtDNA is generally more useful than nuclear DNA in tracing genetic relationships in historical contexts. With one exception, nuclear DNA is at present useless for genealogical research over a wide time-gap, because there is currently no way of determining which components of the nuclear DNA are derived from which ancestor. In fact, many ancestors may be represented by no nuclear DNA components in their living descendants. The one certain exception to this rule is the Y-chromosome – and we shall return to this point later, because it is of potential interest in the attempt to establish an overall picture of the DNA of the Yorkist princes.

For the moment, however, let us consider only mitochondrial DNA. Occasional spontaneous mutations occur in mtDNA, and these are then passed on to descendants, though they may take up to six generations to become firmly established. Such mutation occurs on average once every 10,000 years. Thus it is possible, by comparing the mitochondrial DNA of two individuals, to establish roughly how much time has elapsed since the lifetime of their last common ancestress in the female line. It has been calculated that all human beings now living are descended in the exclusively female line from one single woman, known as Mitochondrial Eve, who lived in Africa about 150,000 years ago. It is argued that every human being now living on the planet can trace his or her mitochondrial DNA back to Mitochondrial Eve. Of course, the latter was not the only living woman of her time and place. What is unique about her is the fact that she is the only one of her contemporaries to have living descendants in the female line.

It is likewise posited that most of the historic native population of Europe can trace their female line ancestry back to one of only seven ‘clan mothers’ who lived between 45,000 and 10,000 years ago. Each of these clan mothers was a descendant of Mitochondrial Eve. The seven clan mothers of Europe are usually referred to by letters, or names, as follows:²¹

U (‘Ursula’) – ancestress of about eleven per cent of the European population. She probably lived in Greece about 45,000 years ago. Her descendants are especially to be found in western Britain.

X (‘Xenia’) – probably lived in Russia about 25,000 years ago. Her descendants (about six per cent of the population) are to be found today mostly in central and Eastern Europe.

H (‘Helena’) – lived in the Bordeaux region of France about 20,000 years ago. Hers is the most widespread European clan, with about forty-seven per cent of the modern population descending from her in the female line.

V (‘Velda’) – probably lived 17,000 years ago in northern Spain, near Santander. About five per cent of native Europeans belong to this clan, which is found mainly in Western Europe.

T (‘Tara’) was more or less a contemporary of Velda. She probably lived in Tuscany. Her descendants, who account for about nine per cent of the modern population of Europe, live mostly along the Mediterranean coast or the western edge of the continent, including western Britain and Ireland.

K (‘Katrine’) – probably lived 15,000 years ago, in the Venice region. She is the clan mother of six per cent of modern Europeans who are most likely to be found around the Mediterranean. ‘Ötzi’ the ‘iceman’ was one of her descendants.

J (‘Jasmine’) – thought to have lived in Syria about 10,000 years ago. Her people were the ones who introduced farming to Europe. Her descendants are found today either in Spain, Portugal and western Britain, or in central Europe. They seem to represent about seventeen per cent of the European native population.

When DNA is being used to attempt to identify long-dead bones, the first thing to note is that it cannot prove the identity of an individual. Mitochondrial DNA has to be compared with a sample from a known relative, as was done recently in the case of the bones thought to be those of the Russian Imperial family. A mismatch proves for certain that the bones cannot be the person sought, but a match does not prove identity, merely that the bones are those of a person with similar mitochondrial DNA to – and thus a relative in some degree of – the person being searched for. Depending on factors such as how widespread the resulting mitochondrial DNA sequence is in the modern European population, and on the precise set of mutations present, that information will be of greater or lesser significance.

However, the final decision about the identity of archaeologically recovered remains will also depend on a variety of other factors: such as location, age at death, the era from which the remains date, and other evidence suggesting identity. Thus, for example, in the case of the Russian Imperial bones, evidence of ages at death, the location of the remains, how the individuals had died, and how their bodies had been treated after death, all supported the identification of the bones as Romanov remains, in addition to the DNA evidence from multiple sources, which confirmed their possible identification.

FAMILY TREE 2: The female line of descent from Catherine de Roët to Joy Brown (Ibsen).

The mitochondrial DNA of Margaret of York and of all her siblings, including Richard III, was that of her mother, Cecily Neville. Cecily received it from her mother, Joan Beaufort, who in turn had received it from her mother, Catherine de Roët. Rather unfortunately, the beautiful Catherine is very widely referred to by her first husband’s surname, ‘Catherine Swynford’. Genealogically, however, it is always preferable to refer to women by their birth surnames, and in the present instance this is particularly essential, since Richard III and his family were not the descendants of Sir Hugh Swynford, but of Catherine de Roët’s second husband, John of Gaunt. For our purposes it is Catherine’s birth family which is important.

Catherine’s mitochondrial DNA may have come from the mainland of Europe; perhaps from the area we now call Belgium, from the Netherlands, or possibly from Germany or France. Catherine’s father was a knight from Hainault. He is usually named as Sir Payne (or Paon) de Roët (or Roelt), and is often said to have come to England from Hainault with Edward III’s wife, Queen Philippa, of whom he was probably a relative. In fact, his real first name seems to have been Gilles. ‘Paon’ (‘Peacock’?) was merely a nickname.²² Perhaps he liked to look his best. There appears to be no real record surviving of his having served in England. However, he did serve Queen Philippa’s sister, Margaret, the Holy Roman Empress, in Germany, and he also served Edward III as a herald and as Guienne King of Arms in the Aquitaine. Ultimately he died in England, and was buried at old St Paul’s Cathedral.²³

But it was not Catherine’s father, but her unknown mother who was the source of her daughter’s (and thus of Richard III’s) mitochondrial DNA. In modern terms this lady might have been French, German, Belgian or Dutch.²⁴ She seems unlikely to have been English.²⁵ Dame de Roët seems to have borne her husband three daughters, and at least one son. All of these children inherited from their mother the mitochondrial DNA which was in due course to be transmitted by Catherine, the youngest daughter of the de Roët family, to her great-grandson Richard III.

The eldest de Roët daughter, Isabelle (or Elizabeth), seems never to have left her homeland. She entered the convent of the Canonesses at Mons, where she lived, died and was buried. Gilles’ two younger daughters, like their father, lie interred in England. The aristocratic Philippa de Roët attracted the attention of the young poet Geoffrey Chaucer, who was the son of mere merchant stock from Ipswich. ‘It may well have been to highborn, theoretically unreachable Philippa that Chaucer wrote some of his love songs.’²⁶ Chaucer did eventually marry Philippa and had children by her. Philippa died in the late summer or autumn of 1387 and lies buried in the chancel of the church of St Mary the Virgin, East Worldham in Hampshire.²⁷

FAMILY TREE 3: The Chaucer connection.

Her younger sister, Catherine de Roët, was a member of the household of John of Gaunt, where she was employed since at least 1365 as one of the ladies attending on his first wife, Blanche of Lancaster. Soon after entering Blanche’s service, Catherine de Roët married Sir Hugh Swynford of Coleby and Kettlethorpe in Lincolnshire, by whom she had a son, Thomas, and a daughter, Blanche. In 1371 Sir Hugh Swynford died while serving abroad, leaving Catherine a young widow. She was then in her early twenties, and despite his second marriage to Constance of Castile, John of Gaunt soon made Catherine his mistress. Meanwhile, her official position in his household was now that of governess to his children. There have been allegations that Catherine became John’s mistress while her first husband was still alive, but this appears unlikely. Their liaison probably began in 1372, after Hugh Swynford’s death, and Catherine’s first child by John of Gaunt was born in 1373.²⁸ The affair was viewed askance at the time, and Catherine was denounced as a seductress. Nevertheless, the couple’s relationship proved enduring.

After various vicissitudes (and following the death of Constance of Castile), on 13 December 1396 John of Gaunt finally married Catherine in Lincoln Cathedral. This marriage, which made Catherine the second lady in the land, after the queen, caused universal astonishment. It has been described as a mésalliance, but that seems hardly just. Catherine was, after all, probably a relative of John of Gaunt’s mother, the late queen, Philippa of Hainault, and thus a distant cousin of her new husband. One important outcome of their marriage was that Catherine’s illegitimate children by John of Gaunt, all of whom had been given the surname ‘Beaufort’, were legitimised by the Pope in 1396 and by King Richard II in 1397.

Mitochondrial DNA identical to that of Richard III would have been found in all four of Catherine de Roët’s sons: Sir Thomas Swynford, John Beaufort, first Earl of Somerset,²⁹ Henry, Cardinal Beaufort, Bishop of Winchester, and Thomas Beaufort, Duke of Exeter. However, as males none of these was able to pass on this mitochondrial DNA. Nevertheless, in the following generation the same DNA would have been found in all of the numerous Neville and Ferrers children of their sister, Joan Beaufort. Richard III also shared his mitochondrial DNA with the three children of the poet, Geoffrey Chaucer, since Chaucer’s wife Philippa was Catherine de Roët’s sister. Although this relationship with the Chaucers has sometimes been questioned, it was explicitly acknowledged by Cardinal Beaufort, who, in a letter, referred to the poet’s son, Thomas Chaucer, as his cousin.³⁰

Unfortunately, no mitochondrial DNA line of descent from Philippa de Roët (Chaucer) survives to the present day, for Philippa had only one daughter and she became a nun (see Family Tree 3). From Catherine de Roët, Duchess of Lancaster, however, the mtDNA line continues through her daughter, Joan Beaufort, Countess of Westmorland. The latter had many daughters, including Cecily, Duchess of York. I traced female lines of descent from Cecily and all her sisters, producing a huge family tree resembling a spider’s web on my computer. It was necessary to attempt to trace all the possible female lines of descent, because there was no way at the outset of knowing which line (if any) would prove to be continuous down to the present day. In the event, none of the lines from Cecily’s sisters proved to continue to the present.

Of Cecily Neville’s own daughters, two produced children: Anne of York, Duchess of Exeter, and Elizabeth of York, Duchess of Suffolk. At first, Elizabeth looked the most hopeful candidate to have living female line descendants because she had a number of children including several daughters. However, her female lines of descent soon petered out. Anne of York, on the other hand, looked less hopeful initially. She had only two children – both daughters – and the elder of these two had no descendants. Nevertheless, it was the line of Anne of York which was to preserve the mtDNA of Cecily Neville’s children until the present day. I traced an unbroken line of descent, mother to daughter, from Cecily Neville’s eldest daughter, Anne of York, Duchess of Exeter, to Mrs Joy Ibsen in Canada.³¹ This line of descent has a number of interesting features, and in the next chapter we shall follow its history in some detail.