The discovery of the presumably lost grave of the controversial English king Richard III in Leicester (U.K.) was one of the most important archaeological achievements of the last decennium. The skeleton was identified beyond reasonable doubt, mainly by the match of mitochondrial DNA to that of living maternal relatives, along with the specific archaeological context. Since the genetic genealogical analysis only involved the DNA sequences of a single 15th century individual and a few reference persons, biologists might consider this investigation a mere curiosity. This mini-review shows that the unique context of a historical king's DNA also has relevance for biological research per se — in addition to the more obvious historical, societal and educational value. In the first place, the historical identification appeared to be a renewed forensic case realising a conservative statement with statistical power based on genetic and non-genetic data, including discordant elements. Secondly, the observation of historical non-paternity events within Richard III's patrilineage has given rise to new research questions about potential factors influencing the extra-pair paternity rate in humans and the importance of biological relatedness for the legal recognition of a child in the past. Thirdly, the identification of a named and dated skeleton with the known historical context serves as a reference for bioarchaeological investigations and studies on the spatio-temporal distribution of particular genetic variance. Finally, the Richard III case revealed privacy issues for living relatives which appear to be inherent to any publication of genetic genealogical data.

Introduction

The discovery of the presumably lost grave of the English king Richard III (1452–1485) underneath a car park in 2012 caught the attention of a broad international public [1,2]. The fascination was mainly stirred up by Richard III's general fame, the controversy surrounding his popular image as a cruel and powerful person, and the as-yet-mysterious circumstances in which he became king and died 2 years later in battle [3,4]. The chronicle narrating the miraculous finding and the oldest cold case, to date, only increased public interest [5]. The genetic identification analyses to be performed on any interesting skeleton found were already announced before excavations started on the Grey Friars site in Leicester, the birthplace of DNA fingerprinting [1]. They were performed on a male skeleton excavated in the choir of the former church, which was later identified as being Richard III.

Although researchers at that time had already successfully performed historical identifications using DNA, e.g. the Romanov family [6,7], the identification of a 15th century individual was still a huge achievement [8,9]. The first prerequisites needed for this feat were to discover the resting place of the person under study, to receive permission for excavation, and to find enough qualitative human remains to perform molecular analyses. This is no easy task, which the many unfruitful attempts to conclusively identify remains of Richard III's sister Margaret in Mechelen (Belgium) illustrate [2,10]. A second necessity was finding appropriate and consenting relatives to adopt the so-called genetic genealogical approach for identification. These relatives are often several generations removed from the individual in question; therefore, only non-recombining DNA markers might be informative. Thanks to linear inheritance, any biological relative in direct maternal or paternal line carries a closely related mitochondrial DNA (mtDNA) or Y-chromosomal haplotype, respectively [11]. Nevertheless, only a minority of ancestors has such currently living direct descendants or relatives [12]. Finally, ancient DNA handling and analysis require specific expertise. To date, only a limited amount of DNA data from Richard III's remains is available, including the mitogenome and Y-chromosomal profile used in the genetic genealogical approach. Additional DNA analyses have been performed to predict the eye and hair colour in order to realise a facial reconstruction (Figure 1a) [13]. The sequencing of the complete genome of Richard III has been announced, but not yet accomplished [14].

The genetic identification of Richard III's remains resulted in a representative image of this individual and in an official cenotaph for one of England's most famous and controversial kings.

Figure 1.
The genetic identification of Richard III's remains resulted in a representative image of this individual and in an official cenotaph for one of England's most famous and controversial kings.

(a) While no portrait made during his life is known, DNA-based predictions appeared to match this post-mortem portrait of Richard III from the 1510s [13] by kind permission of the Society of Antiquaries of London. (b) The inauguration of this permanent repository of Richard III's remains in Leicester's cathedral marked the closing of a complex and much-discussed genetic identification process [2] (source: author, May 2015).

Figure 1.
The genetic identification of Richard III's remains resulted in a representative image of this individual and in an official cenotaph for one of England's most famous and controversial kings.

(a) While no portrait made during his life is known, DNA-based predictions appeared to match this post-mortem portrait of Richard III from the 1510s [13] by kind permission of the Society of Antiquaries of London. (b) The inauguration of this permanent repository of Richard III's remains in Leicester's cathedral marked the closing of a complex and much-discussed genetic identification process [2] (source: author, May 2015).

The identification of Richard III was immediately put forward as the most important archaeological discovery of the 21st century [15]. The remains did partly reveal the physical appearance of and the real story behind this king, whose famous image was until then mainly formed by controversial Shakespearean literature hailing from the Tudor era [16]. Subsequently, many articles focused on insights relevant for historical and archaeological sciences [2,1719]. Moreover, the discovery of such cultural heritage has also direct socio-economic relevance: it rapidly boosted the international profile of Leicester and its university, which has been appreciated as an invaluable PR stunt [20]. The funeral of the Richard III was a large event that increased cohesion among citizens [21] and brought inspiring challenges for the city's multicultural atmosphere and image [22]. The visitors to the repository of the remains in Leicester cathedral (Figure 1b) leave a substantial and durable economic impact [23]. The identification also encouraged a broad and young public to acquire knowledge in history and science. The Richard III case has even been noted as an educative example among scholars since it clearly demonstrated the importance of multidisciplinary research. Forensic geneticists and pathologists, osteologists, archaeologists, weapon experts, engineers, Latinists, historians, and genealogists worked together and successfully united the fields of science and humanities [20]. However, this case study is not a curiosity with a merely historical, societal, and educational value: here, we particularly focus on the genetic identification of Richard III and its relevance for specific research fields in biology, four of which are discussed below.

Forensic genetic identification and the public

The Richard III investigation clearly showed the complexity and caveats of the genetic genealogical approach within forensic genetics, a discipline that beyond adding genetic data to a biological trace or remain also needs to communicate results in a transparent and conclusive way [24,25]. The mitogenome sequence of the skeleton matched with that of two living maternal relatives whose most recent common ancestor (MRCA) was Richard III's grand-niece (Figure 2). The lineage was neither found among the 1823 samples of a British mtDNA database nor in the 26 127 European haplotypes stored in EMPOP (https://empop.online) [13,26]. Among the >250 000 accessible mtDNA profiles of direct-to-customer (DTC) genetic testing companies, only seven independent hits were found [27]. Therefore, a coincidental match between the skeleton and the reference persons seemed highly unlikely. However, the skeleton's Y-chromosomal lineage differed from that of five paternal relatives of Richard III (Figure 2). The MRCA of the latter was the 5th Duke of Beaufort (1744–1803), whose MRCA with Richard III was Edward III (1312–1377). One of the five revealed a non-paternity event during the last five generations. The remaining four relatives, however, were also assigned to a different lineage than the one attributed to Richard III, meaning the DNA evidence alone did not suffice to assure the identity [13]. Criticism on the statement of identification was even formulated publicly when the different research aspects were considered separately [28]. Therefore, the researchers used an innovative Bayesian statistical approach combining probabilities for all genetic and non-genetic elements. They included facts corresponding between known history and the observed archaeological context, such as radiocarbon dating, sex, age estimation, scoliosis, and perimortem wounds consistent with medieval battle injuries, as well as the discordant elements, such as a low historical extra-pair paternity (EPP) rate to deal with the Y-chromosomal mismatch. The evidence for a positive identification was extremely strong after such integrative analysis [13]. The researchers systematically tested each alternative, even controversial, hypothesis that might explain the results, including the double hypothesis, according to which a male maternal relative of the king was taken on the battlefield [1]. Consequently, the Richard III case is viewed as an example in identification studies (i) for the application of a statistical method combining all variables when DNA data only have a limited value or may include inconsistencies and (ii) for formulating and statistically testing alternative scenarios in contrast with previous studies where only a DNA match with living relatives was declared sufficient evidence [29].

Pedigree showing the genealogical links between king Richard III and the living male-line (given in blue) and female-line (given in green) relatives who participated in the genetic identification study of King et al. [13].

Figure 2.
Pedigree showing the genealogical links between king Richard III and the living male-line (given in blue) and female-line (given in green) relatives who participated in the genetic identification study of King et al. [13].

Numbers indicate the amount of anonymous individuals in the genealogy between named individuals. The individuals given in red were born illegitimate and were later legitimised. Figure adapted from [13,39].

Figure 2.
Pedigree showing the genealogical links between king Richard III and the living male-line (given in blue) and female-line (given in green) relatives who participated in the genetic identification study of King et al. [13].

Numbers indicate the amount of anonymous individuals in the genealogy between named individuals. The individuals given in red were born illegitimate and were later legitimised. Figure adapted from [13,39].

Finally, awareness about the complexity of DNA typing and statistical assessment in forensic cases is raised by broad media coverage of such investigations. This may become crucial as results of genetic identification have to be interpreted correctly also by non-experts who have to take decisions at court or in politics [24,25,30]. Nevertheless, there were also negative reactions by professionals immediately after the press conference on the Richard III case. The researchers in Leicester announced results that were not accessible at that time in order to verify the genetic ‘matches’ [20]. Elsewhere, unjustified but publicly claimed ‘matches’ using the genetic genealogical approach had to be retracted after statistical analysis was found inadequate (e.g. [29,3133]). Therefore, the Richard III case showed that it is important for the credibility of a discipline to provide in-depth results or a peer-reviewed publication when an identification is claimed publicly.

EPP behaviour

In an EPP event, the social father is (unknowingly) not the biological father of his child [34]. The frequency and factors that influence the EPP rate are highly investigated in many pair-bound species, as males are investing in paternal care without any direct benefit for their own fitness. Ironically, the knowledge on humans is still limited [35]. Genetic genealogical research provides insights by testing potential factors on human EPP behaviour in the past [3638]. The identification of Richard III presented only a single familial line but with the remarkable observation of at least two historical EPP events [13]. Beyond wild speculations on when these occurred and historical ‘gossip’ with likely political motives [39], the observations in this specific patrilineage give rise to biological research questions about human cuckoldry behaviour. One important question is whether the EPP rate was and still is different between socio-economical classes within a population [36]. At the time, it was essential for a royal family to have (male) heirs, illustrated by the political disaster of the death of Richard III's only son during his short reign [3,4]. Since EPP was a rarely raised political issue and the legitimacy of their wives' child could not be opposed legally by others, males in noble families might have accepted EPP to maintain continuity and political stability [39]. This specific example reveals the necessity to investigate differences in the (historical) EPP rates depending on relevant inherited property and political motives [40]. Another important question is raised by the fact that the patrilineage between Richard III and his living relatives included two ancestors who were not legally recognised by a father after birth but later when their mothers married (Figure 2) [13,39]. In patrilineages including premarital children, the chance of an observed EPP event is assumed to be much higher, but data are still lacking [41]. More research on such patrilineages, like the one of Richard III, will provide insights in evolutionary and historical demography by revealing how often and under which circumstances males invested in non-biological children.

Bioarchaeology

Bioarchaeology describes the contextual analysis of biological remains from past societies to realise comparative studies on, for example, violence, colonialism, and health [42]. Remains of a single individual, like that of Richard III, may contribute to these broad studies because the known point of time and life story associated with the remains make it possible to validate specific results of bioarchaeological — including genetic — investigations. This improves the interpretation of (many) other anonymous skeletons of individuals which possibly lived or died under similar circumstances in the same period. Most attempts to discover such ‘identifiable graves’ to use as reference were not successful [43]. The single case of Richard III did already have wide implications for non-genetic methodologies, like isotope-based palaeodietary and migration reconstructions. The isotopic data of the skeleton would suggest that Richard III had migrated to a different area in the last few years of his life [44]. As this was not the case according to known history, the results were explained by dietary differentiation since he became king. A similar validation exercise was realised within tool mark analysis: traumata on the named skeleton could be interpreted using data known from battles, weapons, and armour of that time [19,45,46]. Research also provided insights into food patterns [44], medical care [4749], and hygienic conditions of the highest social class in the Late Middle Age [50], something that was not possible with anonymous graves. Genetic investigations on the micro-level of a known individual will enable spatio-temporal analyses to locate the occurrence of genetic variants within a well-known historical and familial context [51,52]. This might become relevant when the whole genome of Richard III will be available, e.g. to investigate if genetic factors might explain his scoliosis [48].

It is generally accepted by scholars that our characteristics are co-constructed by genetic and environmental factors. Still, deterministic accounts remain popular [53,54], also because of the widely advertised idea that DNA informs our sense of identity and best lifestyle [55,56]. Therefore, the study of Richard III's full genome might become a challenging exercise for bioarchaeologists and geneticists in interpreting and communicating genetic variants for a single individual. Owing to the biographical details and controversial character, this interpretation will be especially thought-provoking in the Richard III case if variants are associated with personality or psychological makeup. Such discussions already appeared when a link between the king's distorted physique and the character was suggested immediately after identification of his skeleton [16].

Genetic privacy

Since the ‘next-generation sequencing revolution’, many initiatives are taken to maintain the privacy of DNA donors when genetic data are publicly available [57]. Anonymous publication is, however, hardly feasible for genetic identifications in which the combination of name and genetic data is essential per se. Nevertheless, any debate about the privacy of an individual that died several centuries ago is almost of a philosophical nature, especially because there seems to be no one that would be harmed by publishing data [58,59]. Still, the Richard III case revealed the difficulty of guaranteeing genetic privacy in the context of the publication of DNA results together with patri- and matrilineages of living relatives, even in a carefully performed historical study [13]. All DNA donors gave a detailed informed consent to analyse and publish their results (Turi King, personal communication), but a privacy issue is still existing for all other family members assumably carrying the same Y-chromosomal or mtDNA lineage. Any person may test their relatedness via a commercial DTC genetic test, with consequences for genetic anonymity [60] and kinship inference [11]. The lack of counselling in those cases is substantial and an unexpected result is a radical event for every party involved [61], not only when a pedigree is the reason for societal privileges [62]. The consequences for families of publishing genetic genealogical information are often only realised afterwards [29,63]. Sequence data might additionally affect privacy when variants are related to medical conditions [64], as already illustrated for Richard III's mitogenome [13].

Excluding genetic genealogical data from publication in an identification case is not an option either. Since the Richard III study, several approaches were proposed to circumvent the privacy issue. An ethical analysis described the (theoretical) possibility of a familial or generational consent in which DNA donors have to inform close family members and all third parties [65]. Another solution was realised in the forensic identification of a blood stain attributed to the Belgian king Albert I, where independent external review of (the quality of) the data and statistical interpretation guaranteed scientific accuracy. The complete methodology, statistical analysis, and the names of the DNA donors were published; however, no DNA information was given to guarantee the genetic privacy of living relatives [66].

Conclusion

Genetic information attributed to a single historical individual might seem of highly restricted biological relevance at first glance. Here, we illustrated that biologists and geneticists might benefit from taking ‘celebrity genetics’ seriously. Historical identification cases trigger new research questions and are an opportunity to validate and communicate results in several biological disciplines.

Abbreviations

     
  • DTC

    direct to customer

  •  
  • EPP

    extra-pair paternity

  •  
  • MRCA

    most recent common ancestor

  •  
  • mtDNA

    mitochondrial DNA

Funding

M.H.D.L. is a postdoctoral fellow of the Fund for Scientific Research — Flanders (FWO-Vlaanderen). M.B. was supported by the Tyrolean Science Fund (Wissenschaftsfonds des Landes Tirol) [UNI-404/1998]. Funding was provided by KU Leuven [BOF-C1 grant C12/15/013] and the Fund for Scientific Research — Flanders [research grant number 1503216N].

Acknowledgments

The authors thank the Editorial committee of Biochemical Society Transactions for the invitation to write this short review. The authors especially thank Turi King and Walther Parson for inspiring discussions, as well as Sarah Princen and two anonymous reviewers for useful comments on a previous version of this manuscript.

Competing Interests

The Authors declare that there are no competing interests associated with the manuscript.

References

References
1
Morris
,
M.
and
Buckley
,
R.
(
2013
)
Richard III — The King Under the Car Park
,
University of Leicester
,
Leicester
2
Carson,
A.J
,
Ashdown-Hill,
J.
,
Johnson,
D.
,
Langley,
P.J.
and
Johnson,
W.
(
2017
)
Finding Richard III: The Official Account of Research by the Retrieval and Reburial Project
,
Troubador Publishing Ltd
,
Leicester
,
200
p
3
Ross
,
C.
(
1981
)
Richard III
,
Eyre Methuen
,
London
4
Carson
,
A.
(
2009
)
Richard III: The Maligned King
,
The History Press
,
London
5
Pitts
,
M.
(
2014
)
Digging for Richard III: The Search for the Lost King
,
Thames and Hudson
,
London
6
Gill
,
P.
,
Ivanov
,
P.L.
,
Kimpton
,
C.
,
Piercy
,
R.
,
Benson
,
N.
,
Tully
,
G.
et al. 
(
1994
)
Identification of the remains of the Romanov family by DNA analysis
.
Nat. Genet.
6
,
130
135
7
Coble
,
M.D.
,
Loreille
,
O.M.
,
Wadhams
,
M.J.
,
Edson
,
S.M.
,
Maynard
,
K.
,
Meyer
,
C.E.
et al. 
(
2009
)
Mystery solved: the identification of the two missing Romanov children using DNA analysis
.
PLoS ONE
4
,
e4838
8
Nilsson
,
M.
,
Possnert
,
G.
,
Edlund
,
H.
,
Budowle
,
B.
,
Kjellström
,
A.
and
Allen
,
M.
(
2010
)
Analysis of the putative remains of a European patron Saint–St. Birgitta
.
PLoS ONE
5
,
e8986
9
Bauer
,
C.M.
,
Bodner
,
M.
,
Niederstätter
,
H.
,
Niederwieser
,
D.
,
Huber
,
G.
,
Hatzer-Grubwieser
,
P.
et al. 
(
2013
)
Molecular genetic investigations on Austria's patron saint Leopold III
.
Forensic Sci. Int. Genet.
7
,
313
315
10
Cassiman
,
J.J.
and
Raeymaekers
,
P.
(
2009
)
Missie DNA: over verwantschap en vreemdgaan, mysteries en misdaad, evolutie en gezondheid
,
Davidsfonds
,
Leuven
11
Calafell
,
F.
and
Larmuseau
,
M.H.D.
(
2017
)
The Y chromosome as the most popular marker in genetic genealogy benefits interdisciplinary research
.
Hum. Genet.
136
,
559
573
12
Helgason
,
A.
,
Hrafnkelsson
,
B.
,
Gulcher
,
J.R.
,
Ward
,
R.
and
Stefánsson
,
K.
(
2003
)
A populationwide coalescent analysis of Icelandic matrilineal and patrilineal genealogies: evidence for a faster evolutionary rate of mtDNA lineages than Y chromosomes
.
Am. J. Hum. Genet.
72
,
1370
1388
13
King
,
T.E.
,
Fortes
,
G.G.
,
Balaresque
,
P.
,
Thomas
,
M.G.
,
Balding
,
D.
,
Maisano Delser
,
P.
et al. 
(
2014
)
Identification of the remains of King Richard III
.
Nat. Commun.
5
,
5631
14
Pappas
,
S.
(
2014
)
King Richard III's Genome to be Sequenced
,
LiveScience
15
Vai
,
S.
,
Lari
,
M.
and
Caramelli
,
D.
(
2016
)
DNA sequencing in cultural heritage
.
Topics Curr. Chem.
374
,
8
16
Kostihova
,
M.
(
2016
)
Digging for perfection: discourse of deformity in Richard III's excavation
.
Palgrave Commun.
2
,
16046
17
Buckley
,
R.
,
Morris
,
M.
,
Appleby
,
J.
,
King
,
T.
,
O'Sullivan
,
D.
and
Foxhall
,
L.
(
2013
)
‘The king in the car park’: new light on the death and burial of Richard III in the Grey Friars church, Leicester, in 1485
.
Antiquity
87
,
519
538
18
Licence,
A.
(
2014
)
Richard III: The Road to Leicester
,
Amberley Publishing
,
Stroud, U.K.
,
96
p
19
Appleby
,
J.
,
Rutty
,
G.N.
,
Hainsworth
,
S.V.
,
Woosnam-Savage
,
R.C.
,
Morgan
,
B.
,
Brough
,
A.
et al. 
(
2015
)
Perimortem trauma in King Richard III: a skeletal analysis
.
Lancet
385
,
253
259
20
Mirza
,
A.
(
2015
)
The king under the car park
.
Perspectives Policy Pract. Higher Educ.
19
,
28
32
21
Halse
,
M.
(
2017
)
Planning for the Leicester City Football Club victory parade
.
J. Bus. Continuity Emergency Planning
10
,
217
229
PMID:
[PubMed]
22
Hassen,
I.
and
Giovanardi,
M.
The difference of ‘being diverse’: City branding and multiculturalism in the ‘Leicester Model’
.
Cities
.
In press
23
Shellard,
D.
(
2016
) A king rediscovered: The economic impact of Richard III and Richard III on the City of Leicester. In
Shakespeare's Cultural Capital
(
Shellard
,
D.
and
Keenan
,
S.
, eds).
Palgrave Macmillan
,
London
24
Larmuseau
,
M.H.D.
,
Cassiman
,
J.-J.
and
Decorte
,
R.
(
2014
)
Controversial identification in a historical case is illustrative of the complexity of DNA typing in forensic research. Response to Charlier et al
.
Forensic Sci. Int. Genet.
9
,
e18
e19
25
Butler,
J.M
. (
2012
)
Advanced Topics in Forensic DNA Typing: Methodology
,
Elsevier Inc.
,
London
,
680
p
26
Parson
,
W.
and
Dür
,
A.
(
2007
)
EMPOP — a forensic mtDNA database
.
Forensic Sci. Int. Genet.
1
,
88
92
27
Logan
,
I.S.
and
Brinkman
,
D.N.
(
2017
)
King Richard III and his mitochondrial DNA haplogroup J1c2c3
.
J. Genealogy Fam. Hist.
1
,
1
14
28
Selwood,
D.
(
2015
) Richard III: we're burying the wrong body.
Telegraph
29
Larmuseau
,
M.H.D.
,
Delorme
,
P.
,
Germain
,
P.
,
Vanderheyden
,
N.
,
Gilissen
,
A.
,
Van Geystelen
,
A.
et al. 
(
2014
)
Genetic genealogy reveals true Y haplogroup of House of Bourbon contradicting recent identification of the presumed remains of two French Kings
.
Eur. J. Hum. Genet.
22
,
681
687
30
Larmuseau
,
M.H.D.
,
Nivelle
,
K.
and
Decorte
,
R.
(
2017
)
Jay D. Aronson (2016) Who owns the dead? The science and politics of death at Ground Zero
.
Forensic Sci. Int. Genet.
28
,
e53
31
Charlier
,
P.
,
Olalde
,
I.
,
Solé
,
N.
,
Ramírez
,
O.
,
Babelon
,
J.-P.
,
Galland
,
B.
et al. 
(
2013
)
Genetic comparison of the head of Henri IV and the presumptive blood from Louis XVI (both Kings of France)
.
Forensic Sci. Int. Genet.
226
,
38
40
32
Olalde
,
I.
,
Sánchez-Quinto
,
F.
,
Datta
,
D.
,
Marigorta
,
U.M.
,
Chiang
,
C.W.K.
,
Rodríguez
,
J.A.
et al. 
(
2015
)
Genomic analysis of the blood attributed to Louis XVI (1754–1793), king of France
.
Sci. Rep.
4
,
4666
33
Delorme
,
P.
(
2013
)
La mauvaise tête de Henri IV, F. Aimard et Y. Briend editeurs, Paris, 384 p (ISBN 978-2-36918-004-3)
34
Gray,
P.B.
and
Anderson,
K.G
. (
2010
)
Fatherhood — Evolution and Human Paternal Behavior
,
Harvard University Press
,
Cambridge, U.S.A.
,
304
p
35
Anderson
,
K.G.
(
2006
)
How well does paternity confidence match actual paternity? Evidence from worldwide nonpaternity rates
.
Curr. Anthropol.
47
,
513
520
36
Larmuseau
,
M.H.D.
,
Matthijs
,
K.
and
Wenseleers
,
T.
(
2016
)
Cuckolded fathers rare in human populations
.
Trends Ecol. Evol.
31
,
327
329
37
Larmuseau
,
M.H.D.
,
Matthijs
,
K.
and
Wenseleers
,
T.
(
2016
)
Long-term trends in human extra-pair paternity: increased infidelity or adaptive strategy? A reply to Harris (2016)
.
Trends Ecol. Evol.
31
,
663
665
38
Strassmann
,
B.I.
,
Kurapati
,
N.T.
,
Hug
,
B.F.
,
Burke
,
E.E.
,
Gillespie
,
B.W.
,
Karafet
,
T.M.
et al. 
(
2012
)
Religion as a means to assure paternity
.
Proc. Natl Acad. Sci. U.S.A.
109
,
9781
9785
39
Ormrod
,
W.M.
(
2016
)
The DNA of Richard III: false paternity and the royal succession in later medieval England
.
Nottingham Medieval Studies
60
,
187
226
40
Larmuseau
,
M.H.D.
,
Claerhout
,
S.
,
Gruyters
,
L.
,
Nivelle
,
K.
,
Vandenbosch
,
M.
,
Peeters
,
A.
et al. 
(
2017
)
Genetic-genealogy approach reveals low rate of extra-pair paternity in historical Dutch populations
.
Am. J. Hum. Biol.
29
,
e23046
41
Larmuseau,
M.HD
. (
2016
) Genetic genealogy 2.0: verifying biological relatedness in historical demographic data. In
The Future of Historical Demography
(
Matthijs
,
K.
,
Hin
,
S.
,
Kok
,
J.
and
Matsuo
,
H.
, eds), pp.
85
87
.
Acco
,
Leuven
42
Stojanowski
,
C.M.
and
Duncan
,
W.N.
(
2015
)
Engaging bodies in the public imagination: bioarchaeology as social science, science and humanities
.
Am. J. Hum. Biol.
27
,
51
60
43
Lehouck
,
A.
,
Van Acker
,
J.
,
Vanclooster
,
D.
,
Decorte
,
R.
,
Gonissen
,
J.
,
Larmuseau
,
M.H.D.
et al. 
(
2016
)
Het schrijn van de Zalige Idesbald in de O.L.V.-ter-Potterie: wie ligt er in de kist? (Koksijde-Brugge, W-Vl)
.
Archaeologia Mediaevalis
39
,
93
96
44
Lamb
,
A.L.
,
Evans
,
J.E.
,
Buckley
,
R.
and
Appleby
,
J.
(
2014
)
Multi-isotope analysis demonstrates significant lifestyle changes in King Richard III
.
J. Archaeol. Sci.
50
,
559
565
45
Brough
,
A.
,
Morgan
,
B.
,
Robinson
,
C.
,
Appleby
,
J.
,
Buckley
,
R.
and
Rutty
,
G.
(
2016
)
Biological profiling of Richard III using post-mortem computed tomography scanning
.
J. Forensic Radiol. Imaging
5
,
31
37
46
Bonney
,
H.E.
(
2015
)
Richard III: skeletal evidence of perimortem trauma
.
Lancet
385
,
210
47
Rai
,
A.
(
2013
)
Richard III — the final act
.
Br. Dent. J.
214
,
415
417
48
Lund
,
M.A.
(
2015
)
Richard's back: death, scoliosis and myth making
.
Med. Humanit.
41
,
89
94
49
Appleby
,
J.
,
Mitchell
,
P.D.
,
Robinson
,
C.
,
Brough
,
A.
,
Rutty
,
G.
,
Harris
,
R.A.
et al. 
(
2014
)
The scoliosis of Richard III, last Plantagenet King of England: diagnosis and clinical significance
.
Lancet
383
,
1944
50
Mitchell
,
P.D.
,
Yeh
,
H.-Y.
,
Appleby
,
J.
and
Buckley
,
R.
(
2013
)
The intestinal parasites of King Richard III
.
Lancet
382
,
888
51
van der Zwaag
,
P.A.
,
van Rijsingen
,
I.A.W.
,
de Ruiter
,
R.
,
Nannenberg
,
E.A.
,
Groeneweg
,
J.A.
,
Post
,
J.G.
et al. 
(
2013
)
Recurrent and founder mutations in the Netherlands — phospholamban p.Arg14del mutation causes arrhythmogenic cardiomyopathy
.
Neth. Heart J.
21
,
286
293
52
Claes
,
G.R.F.
,
van Tienen
,
F.H.J.
,
Lindsey
,
P.
,
Krapels
,
I.P.C.
,
Helderman-van den Enden
,
A.T.J.M.
,
Hoos
,
M.B.
et al. 
(
2016
)
Hypertrophic remodelling in cardiac regulatory myosin light chain (MYL2) founder mutation carriers
.
Eur. Heart J.
37
,
1815
1822
53
Moore
,
D.S.
(
2008
)
Espousing interactions and fielding reactions: addressing laypeople's beliefs about genetic determinism
.
Philos. Psychol.
21
,
331
348
54
Radick
,
G.
(
2016
)
Teach students the biology of their time
.
Nature
533
,
293
55
Su
,
P.
(
2013
)
Direct-to-consumer genetic testing: a comprehensive view
.
Yale J. Biol. Med.
86
,
359
365
PMID:
[PubMed]
56
Phillips
,
A.D.
(
2016
)
Only a click away — DTC genetics for ancestry, health, love…and more: a view of the business and regulatory landscape’
.
Appl. Transl. Genomics
8
,
16
22
57
Erlich
,
Y.
and
Narayanan
,
A.
(
2014
)
Routes for breaching and protecting genetic privacy
.
Nat. Rev. Genet.
15
,
409
421
58
Appel,
J.M.
(
2012
)
Privacy versus history: how far should the dead hand reach
?
Camb. Q. Healthc. Ethics
21
,
51
63
59
Holm
,
S.
(
2001
)
The privacy of Tutankhamen — utilising the genetic information in stored tissue samples
.
Theor. Med.
22
,
437
449
60
Borry
,
P.
,
Rusu
,
O.
,
Dondorp
,
W.
,
De Wert
,
G.
,
Knoppers
,
B.M.
and
Howard
,
H.C.
(
2014
)
Anonymity 2.0: direct-to-consumer genetic testing and donor conception
.
Fertil. Steril.
101
,
630
632
61
Moray
,
N.
,
Pink
,
K.E.
,
Borry
,
P.
and
Larmuseau
,
M.H.D.
(
2017
)
Paternity testing under the cloak of recreational genetics
.
Eur. J. Hum. Genet.
25
,
768
770
62
Bowcott
,
O.
(
2016
)
DNA Evidence Proves Accountant is True Heir to Scottish Baronetcy
,
The Guardian
63
Williams
,
S.R.
(
2005
)
Genetic genealogy: the Woodson family's experience
.
Cult. Med. Psychiatry
29
,
225
252
64
Peltzer
,
A.
,
Mittnik
,
A.
,
Wang
,
C.-C.
,
Begg
,
T.
,
Posth
,
C.
,
Nieselt
,
K.
et al. 
(
2018
)
Inferring genetic origins and phenotypic traits of George Bähr, the architect of the Dresden Frauenkirche
.
Sci. Rep.
8
,
2115
65
Wallace,
S.E.
,
Gourna,
E.G.
,
Nikolova,
V.
and
Sheehan,
N.A.
(
2015
)
Family tree and ancestry inference: is there a need for a ‘generational’ consent?
BMC Med. Ethics
16
,
87
66
Larmuseau
,
M.H.D.
,
Bekaert
,
B.
,
Baumers
,
M.
,
Wenseleers
,
T.
,
Deforce
,
D.
,
Borry
,
P.
et al. 
(
2016
)
Biohistorical materials and contemporary privacy concerns - the forensic case of King Albert I
.
Forensic Sci. Int. Genet.
24
,
202
210