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Persian
Pedigree Matters
Earliest Mares in the General
Stud Book
Equine
Genetic Genealogy
Mapping the Thoroughbred
in the Equine Mitochondrial Tree
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Deep Rooted Anomalies in Female Families Revealed by mtDNA
Testing.
Mitochondrial DNA (mtDNA) is passed, with very
rare exception, only from mothers to their offspring. The actual
sequence of nucleotides changes very slowly over time because of
spontaneous mutations that do not interfere with the normal
functions. These two characteristics (inheritance only from the
mother, & gradual creation of diverse sequences over time) make
mtDNA testing potentially useful in identifying & validating
Thoroughbred (TB) female families defined from pedigrees.
That mtDNA typing revealed multiple maternal lines of descent
(matrilineages) in some Thoroughbred families (as defined from
pedigrees) was first reported by Hill, et al (2002). The authors
classified these discrepant lineages as modern, possible de novo
mutation, and deep rooted anomalies. Deep rooted anomalies were
identified in families numbered (in the Bruce Lowe system) 5, 6,
9 and confirmed by Bower, et al (2012b). An additional anomalous
lineage in family 12 was reported by Bower, et al (2012a).
Bower, et al (2012b) also concluded that, despite the
discrepancies reported, mtDNA test results were relatively
consistent within branches (or sub-lineages) of families. Their
proposed explanation for this phenomenon is that "errors in
pedigrees must have occurred largely, though not exclusively, at
sub-lineage foundation events, probably due to incomplete
understanding of modes of inheritance in the past, where
maternal sub-lineages were founded from individuals, related,
but not by female descent."
Recent mtDNA testing of 2000
Thoroughbreds using methods that define the haplotypes more
accurately, linked with detailed pedigree information (Rogers,
2017), has confirmed the presence of the deep-rooted and modern
anomalies identified by Hill & Bower, but is contradictory of
Bower's conclusions regarding when and why these "errors"
largely occurred.
A more plausible explanation for the
reported deep-rooted anomalies is suggested by the historical record. It
should be noted that when Bruce Lowe numbered mare lines
according to their production of winners of the English
classics, he used as his pedigree authority the fifth edition
(1891) of volume one of the General Stud Book (GSB). Three of
these four deep-rooted anomalies can be traced to editorial
changes made in the 5th edition where family lines were linked
that had been separate in earlier editions. The fourth
deep-rooted anomaly probably originated in the early practice of
recording only the names of the stallions in pedigrees. Faced
with two identical sequences of named sires, it was practically
inevitable that later pedigree collectors would consolidate
different matrilineages into the same family.
Changes in
GSB
The 4th edition of Volume One of GSB in 1858
presented pedigrees collected and initially published in nine
preceding editions, the earliest (An Introduction to a General
Stud-book) in 1791. Few substantive changes to the roster of
eighteenth century mares, beyond the addition of a section for
Irish Mares, occurred after the 1808 edition, until 1891. In the
1891 edition, possibly reflecting increased interest in female
lines of descent, as evidenced by the publications of
Stamm-mütter des Englischen Vollblutpferdes by Hermann Goos in
1885 and Familien Tafeln des Englischen Vollbluts, by J D
Frentzel in 1889, the earliest mare with a produce record in
each "direct female line" was identified with a list of her
descendants. Also added was an "index to principal stallions in
the Stud Book up to the year 1890, shewing the original mares
from which they descend in female line." Affirming this
interest, another major editorial change was the presentation of
a separate section devoted to the "earliest known mares, very
few of which have been printed before as brood mares (though the
substance has been given in the Miscellaneous Pedigrees at the
end of earlier editions)…" Lists of descendants are also given
for the "earliest recorded mares of their strains" whose entries
occur in the "second alphabet" or in the addenda. Also added as
an example was a "Table shewing descent in the female line from
an original mare." This shows three branches of descent from
"1st dam" Burton Barb mare to her descendants, Manilla (gr gr
grandam of Blacklock) in the 12th generation, Electress (grandam
of Teddington) in the 15th generation, Martha Lynn (dam of
Voltigeur) and Peri (dam of Sir Hercules) in the 16th
generation, and Bribery (dam of St Albans) in the 17th
generation.
Families with deep-rooted
anomalies
This table shows the affected Bruce Lowe figure
families with the founder mare of each as defined by pedigrees
in the 5th edition. Also shown is the name of the most recent
common ancestor (MRCA) for those sharing a given haplotype and
descent by pedigree. The standard nomenclature for mtDNA typing
is followed here (Achilli, et al, 2012) and when necessary,
other designations converted. |
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| Family ("Founder Mare") |
Haplotype 1 (# tested) |
MRCA |
Haplotype 2 (# tested) |
MRCA |
| 5 (Massey Mare) |
B1a (29) |
Ebony (1728) |
D1b (19) |
Miss West (1777) |
| 6 (Old Bald Peg) |
L2a2 (4) |
Old Morocco Mare |
M2 (17) |
Horatia (1758) |
| 9 (Old Vintner Mare) |
G2a (117) |
Curwen Spot
Mare |
L1a (41) |
sister to
Sloven (1728) |
| 12 (Old Montague Mare) |
I2a2a (8) |
Mother Western (1731) |
L1a (14) |
sister to Sampson (1723) |
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Family 5 (Massey Mare)
29 descendants
(according to GSB 1891 pedigrees) of Ebony (1728) have
haplotype B1a while 19 descendants of Miss West (1777) have D1b.
The explanation appears to be that in the 5th edition, Miss West
was made a descendant of the Massey Mare, whereas her pedigree
in earlier editions had ended with an anonymous Basto mare.
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| GSB, i, 4th ed, 1858 |
GSB, i, 5th ed, 1891 |
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[5]
(OLD) EBONY (black) - sister to Brown Betty,
Bred by the Duke of Rutland, in 1714, got by Basto, out
of his Massey Mare, by Mr Massey's Black Barb. |
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[5]
❖ EBONY (or Ebony Mare),
Foaled in
1728, got by Flying Childers, her dam, Old Ebony, by
Basto. |
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[5]
❖ YOUNG EBONY,
Bred by Lord Portmore,
in 1742, got by Crab, out of the preceding mare [EBONY
(1727)]. |
REGULUS MARE,
Bred by Mr Panton, her dam by Crab
- Childers - Basto. |
[5]
REGULUS MARE (sister to Favourite),
Bred
by Mr Panton, her dam, Y Ebony, by Crab - Ebony, by
Childers - Basto. |
MISS WEST,
Bred by Mr Gee, in 1777, got by
Matchem - Regulus - Crab - Childers - Basto. Died in
1802. |
[5]
MISS WEST,
Bred by Mr Gee, in 1777, got by
Matchem - Regulus - Y Ebony, by Crab - Childers - Basto. |
❖ indicates new or substantially
revised entries in the 1891 edition of GSB |
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Miss West & her dam (a mare by Regulus) have had broodmare
entries in GSB since 1791. Their pedigrees were given through
the 1858 edition as:
MISS WEST,
Bred by Mr GEE, in
1777, got by Matchem - Regulus - Crab - Childers - Basto.
REGULUS MARE,
Bred by Mr PANTON, her dam by Crab - Childers -
Basto.
In the 1891 edition, this lineage was revised to
identify the Crab cross as Young Ebony, & the Childers cross as
Ebony. An undated chesnut filly by Regulus attributed to Mr
Panton was added to the produce of YOUNG EBONY, by Crab. There
is no reference to any new records having come to light to
justify the merging of these two lines. Instead, this would seem
to be an example of what we see even today, of popular stallions
being used in the same succession on mares descending from
different founder mares.
Childers and Basto were
stallions in the Duke of Devonshire's Stud at Chatsworth.
Reference to the list of the Duke of Devonshire's horses at
Chatsworth, 20 Jun 1729 reveals four mares sired by Basto, as
well as numerous currently-unidentified mares (see C M Prior,
The Royal Studs of the Sixteenth and Seventeenth Centuries,
1935; p 123-124). The known Basto daughters were Ebony (and foal
by Childers), Brown Basto Mare (and foal by Childers), a mare by
Basto out of Desdemona, and a mare by Basto, out of Ramsden. GSB
itself includes three mares by Basto with produce by Childers:
(1) BASTO MARE (sister to Soreheels), dam of seven foals by
Childers, including two fillies Bay Basto and Brown Basto;
(2) (OLD) EBONY (sister to Brown Betty), dam of four known
foals, including Ebony, by Childers; and
(3) a mare by
Basto, out of MAKELESS MARE - her daughter by Childers was dam
of Gaul'em, by Starling.
Clearly there were candidates
other than Young Ebony - Ebony that would have "fit" the
sires-only version of Miss West's pedigree. In light of current
mtDNA results, the particular Basto mare in Miss West's line is
best regarded as unidentified, although she may very well also
have come from the Duke of Devonshire's Stud. |
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Family 6 (Old Bald Peg)
4 descendants of
the Old Morocco Mare (according to GSB 1891 pedigrees) have
haplotype L2a2 while 17 descendants of Horatia (1758) have M2.
The explanation appears to be that in the 5th edition, the
Spanker mare at the root of Horatia’s pedigree was assumed to be
the same as the Spanker mare descended from the Old Morocco
Mare. |
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| GSB, i, 4th ed,
1858 |
GSB, i 5th ed,
1891 |
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[6]
OLD
MOROCCO MARE (sometimes called Old Peg),
Bred at Helmsley by Lord FAIRFAX, got by his Morocco
Barb, out of Old Bald Peg, who was got by an Arabian,
out of a Barb mare. |
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[6]
SPANKER MARE,
Bred by LORD DARCY, about 1690,
out of The Old Morocco Mare.
* f Betty Percival, by
[Leedes' Arabian] - Mr Leedes
* BETTY PERCIVAL had a
f by The Paget Turk. |
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MISS BELVOIR,
Bred by the Duke of RUTLAND, got by Grey Grantham –
Paget Turk – Betty Percival, by Leedes’s Arabian –
Spanker. |
[6]
MISS
BELVOIR,
Bred by the Duke of RUTLAND, got by Grey
Grantham - Paget (or Pigot) Turk - Betty Percival, by
Leedes's Arabian - Spanker. |
CHILDERS MARE,
Bred by the Duke of DEVONSHIRE, out of Miss Belvoir, by
Grantham – Paget Turk – Betty Percival, by Leedes’s
Arabian – Spanker. |
[6]
CHILDERS
MARE (sister to Y Miss Belvoir),
Bred by the Duke of
DEVONSHIRE, out of Miss Belvoir, by Grantham - Paget or
Pigot Turk - Betty Percival, by Leedes's Arabian -
Spanker. |
HORATIA (also
called Favourite)
Bred by Mr PANTON, in 1758, got by
Blank, her dam by Childers – Miss Belvoir, by Grantham –
Paget Turk, &c. |
[6-b]
HORATIA
(also called Favourite),
Bred by Mr PANTON, in 1758,
got by Blank, her dam by Childers - Miss Belvoir, by
Grantham - Paget Turk, etc. |
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In editions before 1891, the founder mare of Horatia's line was
an anonymous mare by Spanker, dam of Betty Percival, by the
Leedes' Arabian. In 1891, Betty Percival was added to the
produce of the Spanker Mare, dam of the LEEDES ARABIAN MARE
(sister to Leeds, sometimes called Cream Cheeks). This
assignment by the editor(s) of GSB is not supported by numerous
surviving 18th century pedigrees which never extended Betty
Percival’s pedigree any earlier than her dam. Current mtDNA
results do not support the linkage introduced in 1891 & provide
evidence that the Spanker Mare (dam of Betty Percival) was not
the same as the SPANKER MARE (dam of Cream Cheeks). |
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Family 9 (Old Vintner Mare)
117
descendants of the Curwen Spot mare (according to GSB 1891
pedigrees) have haplotype G2a while 41 descendants of the sister
to the Bolton Sloven have haplotype L1a.
In this case,
from the very earliest editions these two matrilineages have
been considered to be the same. In editions of 1791, 1793, and
1808, the sister to Sloven’s pedigree was given as got by Bay
Bolton - Curwen Bay Barb - Curwen Old Spot - White-legged
Lowther Barb - Vintner Mare.
However, examining pedigrees
that pre-date GSB provides evidence that distinguishes these two
lineages. The Old Vintner Mare was evidently in use in Mr Henry
Curwen’s stud; in fact, a pedigree in Mr Routh’s collection,*
identified her sire as Mr Henry Curwen’s Arabian. Three more
generations judging from the sire’s names were produced in the
Curwen stud. In 1715, Mr Curwen advertised his whole stud for
sale, including "30 Choice Brood Mares, 22 of which Mares are
the Daughters of the famous bay Barbe" (Evening Post, 3 Sep
1715). It may have been at this time, that two sisters to
Mixbury passed into the hands of other breeders. It is unlikely
that all 22 of these daughters of the Curwen Bay Barb were
descended from the Vintner Mare.
___ * C M Prior, Early
Records of the Thoroughbred Horse, 1924.
The sister to
Sloven (MRCA of the second lineage now included in family 9) was
bred by the Duke of Bolton. 18th century pedigrees, notably
those certified by Peter Hammond (identified as Stud-Groom,
1736-1739, and Steward, 1750, to the Duke of Bolton) never
extended her pedigree to the Vintner Mare, ending instead with a
cross of the White-legged Lowther Barb. An advertisement for
Achilles (Newcastle Courant, 11 Jun 1757) is also informative.
In this advertisement, the breeder, William Carr, of Etal,
certified that he had purchased Achilles' dam "of Mr Hammond,
out of the late Duke of Bolton’s Stud."
He gave the
pedigree of his horse as |
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| He was got by the Duke of Bolton’s Little John, his
Dam, by Mogul, Son of the Godolphin Arabian; his
Grand-dam, by Sweepstakes, Son of Lord Oxford’s Bloody
Shoulder’d Arabian; his Great Grand-dam, by the famous
Bay Bolton; his Great Great Grand-dam, by the Curwen Bay
Barb, which got Old Partner’s Dam; his Great Great Great
Grand-dam, by Old Spot, which got Partner’s Grand-dam;
and his Great Great Great Great Grand-dam, by the
Lowther White Legged Barb, which got Partner’s Great
Grand-dam. |
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The pedigree of Achilles' great great grand-dam has the same
sequence of sires as did Partner's dam (sister to Mixbury); that
is Curwen Bay Barb - old Spot - Lowther White Legged Barb.
However, although Carr obviously was pointing out the similarity
in pedigree, he did not make the stronger claim that the mare in
his horse's pedigree was Partner's dam, or a sister to Mixbury.
The Duke of Bolton is known to have owned a "Chesnut
Curwen Mare" (Cheny 1728) whose pedigree was given as by
Curwen's Bay Barb, dam by old Spot. The editor(s) of GSB had no
original records of the early Bolton Stud available, the stud
having been advertised for sale in 1755, after the death of the
third Duke, and again in 1765, after the death of the fifth
Duke. By that time it apparently consisted of only two mares,
mother and daughter, and their produce. In fact, the first two
editions of GSB gave Sloven’s pedigree as by Bay Bolton - Curwen
Bay Barb - Selaby Turk. By the time pedigrees were being
collected for GSB, it is also doubtful that the editor(s) knew
how large the Curwen Stud had been with its potential for
descendants of different founder mares to share the same
sequences of sire names. Given the early practice of providing
pedigrees with only the sires named along the female line of
descent, it was practically inevitable that some separate
matrilineages would have been consolidated into single families
by later pedigree collectors.
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Family 12 (Old Montagu Mare)
8
descendants of the Old Montague Mare (according to GSB 1891
pedigrees) have haplotype I2a2a (as did Eclipse) while 15
descendants of the sister to Sampson (1723) have haplotype L1a.*
In editions predating 1891, the pedigree of sister to Sampson
(GREYHOUND MARE) ended with a Montagu Mare. In 1891, she was
identified with the dam of Mother Western. The conglomeration of
these 2 lines has been questioned in the past based on the
estimated dates of the horses involved. It seems likely that
this was an arbitrary decision based only on the occurrence of
the name Montagu in both pedigrees.
___* This is also the
haplotype found in descendants of the sister to Sloven (1728).
That a line descended from the old Montagu Mare (dam of a
Whiteshirt mare) was present in the Curwen Stud, as well as a
line descended from the Vintner Mare, is evidenced by pedigree
certificates signed by Mr Curwen (see C M Prior's Early Records
of the Thoroughbred Horse, 1924; Mr Cuthbert Routh's Stud-book).
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| GSB, i, 4th ed,
1858 |
GSB, i, 5th ed,
1891 |
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[12-a]
MONTAGU
MARE,
Bred by Lord DARCY, got by his (Old) Montagu,
her dam by Hautboy – Brimmer – Royal Mare.
* f by
Whiteshirt … Mr Smith
* This mare bred a f by Darcy
Chesnut Arabian. |
CURWEN BARB MARE,
Got by the Bay Barb, her dam by the D’Arcy Chesnut
Arabian – Whiteshirt – Montagu Mare. |
[12-a]
❖ CURWEN
BAY BARB MARE (sister to Little George),
Bred by Lord
D’ARCY, got by the Bay Barb, her dam by the D’Arcy
Chesnut Arabian – Whiteshirt – Lord D’Arcy’s Montagu
Mare, by D’Arcy’s Old Montagu – Hautboy – Brimmer. |
GREYHOUND MARE,
Bred by Mr CROFTS, in 1723, out of
Sophonisba’s dam, by the Curwen Bay Barb, page 60. [Got
by the Bay Barb, her dam by the D'Arcy Chesnut Arabian -
Whiteshirt - Montagu Mare]. |
[12-a]
GREYHOUND
MARE (sister to Sampson),
Bred by Mr CROFTS, in 1723,
dam by the Curwen Bay Barb - D'Arcy's Chesnut Arabian. |
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Summary
Three of four deep-rooted
anomalies in Thoroughbred families reported since 2002 can be
traced to editorial changes made in the 5th edition of Volume
One of the General Stud Book. The fourth is probably the result
of confusion pre-dating GSB, arising from the absence of
original records of the studs involved and the early practice of
recording mare lines using only the names of the sires.
© 2018 A J Hibbard |
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References
Achilli A, Olivieri A, Soares P, Lancioni H,
Kashani BH, Perego UA, Nergadze SG, Carossa V, Santagostino M,
Capomaccio S, Felicetti M, Al-Achkar W, Penedo M C,
Verini-Supplize A, Houshmand M, Woodward S R, Semino, O,
Silvestrelli M, Giuliotto E, Pereira L, Bandelt H J, Torroni A:
Mitochondrial genomes from modern horses reveal the major
haplogroups that underwent domestication. PNAS USA. 2012 Feb
14;109(7):2449-54
Hill, EW,
Bradley DG, Al-Barody M, Ertugrul O, Splan RK, Zakharov I,
Cunningham EP. Animal Genetics, 33, 287-294 (2002).
"History and integrity of thoroughbred dam lines revealed in
equine mtDNA variation."
Bower, MA, Campana MG, Nisbet
RER, Weller R, Whitten M, Edwards C J, Stock F, Barrett E,
O'Connell TC, Hill EW, Wilson AM, Howe CJ, Barker G, Binns M.
Archaeometry, first published online: 11 Apr 2012.
"Truth in the bones: resolving the identity of the founding
elite thoroughbred racehorses."
Bower, MA, Whitten M,
Nisbet RER, Spencer M, Dominy KM, Murphy AM, Cassidy R, Barrett
E, Hill EW, Binns M. J Anim Breed Genet (2012) 1-9.
"Thoroughbred racehorse mitochondrial DNA demonstrates closer
than expected links between maternal genetic history and
pedigree records."
Rogers, Byron. CEO, Performance
Genetics. Personal communications. 2017. |
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