Pedigree and Inbreeding Analysis of the Rare and Endangered Akhal-Teke Horse An Honors Thesis (Honors 499) By Jillian Marie Carroll Thesis Advisor Dr. C. Ann Blakey Ball State University Muncie, IN May 2010 Expected Date of Graduation May 2010 Speo)! Y-Ihes/s deC3rQ fJ 0, Lf:' :J.Lj89 , 21./ ABSTRACT ;)O}O • C3 7'7 The Akhal-Teke horse of Turkmenistan is an ancient breed that developed in response to the desertification of Central Asia. The breed has been tightly involved with political upheavals and in 1979 they were introduced into the United States. The Akhal-Teke is currently listed as "Vulnerable" by the Equine Survival Trust and "Threatened" by the American Breed and livestock Conservancy. This may be in part due to the massive inbreeding that is occurring within the population. Inbreeding causes a genetic bottleneck whereby the two alleles of a gene become identical by descent. From a data set of S27 individuals twenty-six pedigrees were computer generated as well as sixteen hand drawn diagrams to investigate the problem and the number of individuals, the relatedness of those horses, and the level of inbreeding were examined in each. Inbreeding coefficients (F) were calculated for specific sire lines in the largest pedigree. Example stallions from each sire line were analyzed and assigned an inbreeding coefficient to demonstrate the effects of unrelated matings on inbreeding. ii DEDICATION To Mr. Joshua Fortriede Without you, these four years would have been very different and I'm fairly sure this thesis may never have had its heart of pedigrees. I am blessed to have you in my life. iii ACKNOWLEDGEMENTS Copious thanks go to Dr. C. Ann Blakey, a wonderful advisor of four years. Your advice, ability to speak some Russian, and flexibility have aided in the writing of this thesis in ways I cannot even express. Heartfelt thanks go to my parents, Mr. and Mrs. John and Susan Carroll, for their everlasting love and support. You two have always been my rock. iv AUTHOR'S STATEMENT The investigation of inbreeding in the Akhal-Teke horse is important because it can act as a mirror project to a similar undertaking with human subjects. There are many more regulations and ethical considerations with human research than with animal research therefore it is much more feasible to begin analysis of a problem as complex and intricate as inbreeding with subjects who can be told to mate with each other without moral consequence. Another reason this project was attainable was the amount of records available. Humans do not necessarily keep track, in a central location, of their family tree. Horse studbooks are exactly that, a centrally located resource that allowed this project to take shape. Whether we would like to admit it or not, inbreeding does occur in our population and the knowledge I have gained during the course of this thesis is knowledge that can be applied to my genetic counseling work in the future. v TABLE OF CONTENTS Page ABSTRACT DEDICATION ii ACKNOWLEDGMENTS iii AUTHOR'S STATEMENT iv LIST OF FIGURES AND TABLES vii CHAPTER 1: literature Review 1 Introduction 1 Background 2 Breed Standards 4 Inbreeding 6 CHAPTER 2: Materials and Methods 9 Materials 9 Methods 9 CHAPTER 3: Results 11 CHAPTER 4: Discussion and Conclusions 17 Discussion 17 vi Conclusions 19 REFERENCES 20 APPENDICES 24 Appendix A: Akhal-Teke Breed Coat Colors 25 Appendix B: Pedigree Data 26 Appendix C: Pedigree Diagram for the Smaller Families 40 Appendix D: Pedigree Diagram for the Large Family 41 vii LIST OF FIGURES AND TABLES Page 13 Figure 1a. Diagrams representing line sires' ancestry for individuals 2a, 44, 165, 9, 213, 100, 432, 697, 828, 26, 569, and 269 . Each dot represents a male individual unless otherwise noted with a «(female). 1\ When read from top to bottom, each line represents a parent-child relationship. refers to the line sire whose lineage is being examined . 14 Figure lb. Diagrams representing line sires' ancestry for individuals 448, 721, 779, and 914. Each dot represents a male individual unless otherwise noted with a «(female). to bottom, each line represents a parent-child relationship. 1\ When read from top refers to the line sire whose lineage is being examined. Table 1. 15 Pedigree data including family information from GenoPro 2007 and information pertaining to related and unrelated individuals. Table 2. 15 Inbreeding calculations (F) for predominant sire line horses based on F* values for recognized ancestral/progenitor sires. Table 3. F values for example stallions from each of the sire lines with an F ~ O. 16 1 CHAPTER 1 Literature Review Introduction The Akhal-Teke horse is an ancient breed originating in Turkmenistan. They were named for the Teke tribe, residents of the Akhal oasis, who prided themselves on their horsemanship. As many Turkmen tribes were nomadic they found a perfect match in the endurance and distance loving Akhal-Teke. Not only could these horses perform in battle but they were good daily workhorses as well (1) . The breed first broke away from its ancestral line of horses roughly 10,000 years ago during the desertification of Central Asia. Due to lack of food and water, the Akhal-Teke horses developed a lighter frame than their ancestors and their high head carriage facilitated survival by boosting the senses of hearing and sight. Sandy camouflage coat color also played into their success in the new desert environment (1). Current world estimates put the total number of breeding mares (females) at just over 900 out of the approximately 4000 known purebred individuals registered with the International studbook in Russia (MAAK) (2). This breed has a restricted studbook and such a small gene pool that the breed is listed as "Vulnerable" on the Equus Survival Trust Equine Conservation List (3), where "Vulnerable" means that there are only 500 to 1500 available breeding mares (females) worldwide. In the United States, the total population of Akhal-Tekes is roughly 400, with only 28 breeding stallions currently listed in the 2009 North American Akhal-Teke Stallion Directory (ATS Directory), provided by Shirley Schulz to the eSTEEDs Project (4). 2 Akhal-Tekes first came to the United States in 1978 with the purchase of a single stallion by Phil and Margot Chase (4). Subsequently a second horse was purchased in 1980 but overall the total purebred population in the United States has remained very limited. The Akhal-Teke Association of America (ATAA) is interested in the propagation and maintenance of the breed and its distinct genetic characteristics through the promotion of the stallions currently available in the 2009 ATS Directory. Breeders can access pedigrees and studbooks to make informed decisions, but may be limited to the registration information, as in the case with the 2009 ATS Directory which only includes information on a maximum of three generations per stallion (4). Therefore, it is possible to misread pedigrees or focus on particular traits without taking into consideration the degree of inbreeding they are propagating. With the lack of detailed pedigree analysis beyond three generations, it is all too easy to make a mistake and breed horses that are only separated by as few as four generations, equivalent to fourth cousin marriages in humans. Background The breed has had a troubled history linked to the rise and fall of various political groups. Before the Bolshevik Revolution, it was not uncommon for every house to have at least one horse. In 1917, the Bolsheviks came into power and revoked the idea of private ownership so all horses were to be rounded up and placed in state run stud farms. Most tribesmen fled with their horses to lands such as Persia and Afghanistan to avoid that fate (1). In 1935, in an attempt to prove the worth of the Akhal-Teke breed, fifteen horses participated in the 4330 kilometer (2600 mile) Ashkhabad-Moscow endurance ride (2). After the 84 day journey, Joseph Stalin consented to the continued breeding of the Akhal-Teke (1) . With the rise of the Soviet Union and the industrialization of agriculture, Khrushchev decided in 1956 that tractors had replaced horses and ordered them to be slaughtered and made into sausage (5). Again the tribesmen came to their beloved breed's 3 rescue and released the horses into the desert. In 1991, breeding became a nationalistic concern and received country-wide attention as an art form . The horse was declared a national treasure and placed on the country's national emblem (1). The political turmoil associated with this breed has continued even into the twenty-first century but this group is not without its well known advocates. One such figure is Geldy Kyarizov, born in 1951 and currently residing in Turkmenistan, immediately took an interest in horses. Though his career took him away from horse breeding, he always had a special place in his heart for the breed. He started a campaign, traveling all of Turkmen istan and the Soviet Union trying to bring recognition to the plight of the breed, culminating the 4330 kilometer ride mentioned previously. This movement peaked public interest which in turn spurred the government to "adopt measures aimed at improving the deplorable situation with horsebreeding in Tu rkmenistan (6)." As part of these new measures in the late 1980s, Geldy established one of the most notable Akhal-Teke breeding farms, named Akhal-Yurt, nestled in the foothills of the Kopetdag mountain range (7). His first well-known stallion was Aigytly (Opal-Agava) and after tasting success he searched Turkmenistan, Southern Kazakhstan and Northern Caucausus for the best studs. Kyarizov's most illustrious acquisition was Yanardag, the horse whose silhouette is on the national emblem. In recognition of his diligent efforts, Geldy was elected the General ,Director of Turkmen Atlary, the Turkmen state equestrian organization (6), On January 30, 2002, Geldy was imprisoned following charges of negligence that are thought to have been staged. His plight became a focal point for the international Akhal-Teke community. One member, Ms. Amrita Ibold, has auctioned off a Limited Edition 2007 Breyer Akhal-Teke model horse three times to aide in Kyarizov's release [auction website : http://www.squidoo.com/geldy 1 (8), He was amnestied by the following president of Turkmenistan in October 2007 (9) . He is still considered to this day to be the Minister of Horses of Turkmenistan . 4 Breed Standards The Akhal-Teke should be a medium-sized horse, giving off the sense of length of body, standing 14.3 to 16 hands tall. The horse should have a rectangular silhouette, strong back, level top line and prominent withers followed by free-moving shoulders. Hips should be wide with a low set tail while the chest is expected to be narrow. A long head, forward set ears, and "hooded" eyes are desirable as is a refined throatlatch as part of a slim neck. Finally, smoothly muscled legs ending in round, hard hooves are essential. Hair ought to be fine covering thin skin with no feathering on the fetlocks . Akhal-Tekes come in many colors and often have white markings (10). A full list of colors and descriptions can be found in Appendix A. Breeding practices in regards to the Akhal-Teke horse can be broken into two schools of thought: the pre-1900 type and the post-1900 type . Turkmen tribes believed in selecting for traits that would benefit them in battle therefore pre-1900 breeding did not focus on body structure. Stamina, speed, and hardiness in the desert were the primary focus of their efforts over physical appearance. As a result of breeding for function over form, two distinct variants developed among the bellicose Turkmen tribes: the warhorses (alaman-at) and the race/state-horses (to i-at) (11). The alaman-at, because of their use in warfare, were more muscular, able to carry heavy loads, and built for long-distance. On the other side of the spectrum the toi-at were more elegant and most often ridden by chiefs of the tribes. Rather than stamina they were created for fast action. Among the more peaceful tribes a third type emerged, the combination horse. They were horses of endurance and finer speed with the ability to haul loads and a sweet character. It bears repeating that through the Turkmen tribes breeding for function, that there were actually three distinct body types produced: alaman-at, toi-at, and the combination body type horse (11). 5 Similar to the formation of distinct body types in the pre-1900s, two "Iooks" arose within the post-1900s: the Russian/Soviet and the Turkmenian, where "Iook" refers to specific sets of refined features . The Russian/Soviet breed form was epitomized by the stallion Boinou (from the toi-at). As a racehorse, he was never tested in the endurance arena yet he was still considered the point of reference for the Russian breeders. By comparison, the Turkmenian breedings were best characterized by the combination of the stallions Everdy-Teleke (from the toi-at) and Karader (from the ala man-at). This era of breeding deviated significantly from the pre-1900s where now form became more important than function. Breeders began to select for particular aesthetic traits such as a "muzzle [that] should fit in a teacup" rather than for general physical strength (12). This approach only served to inhibit function . Because of this appearance-specific focus, the breed began to lose many of its foundation characteristics and center more on the show qualities of that era (12). While written records of Akhal-Teke breeding started the year Boinou was born, 1885, the first studbook was not published until 1941. 287 stallions and 468 mares were included. In 1975 the studbooks were closed when the breed was considered purebred . Post 1973 studbook records have been kept by the All-Russian Institute of Horsebreeding (VNIIK) . Between the years of 1994 and 1997, stallion registrations increased by nearly 40% and mare registrations increased by less than 1%. Registry demands a pedigree and blood sample for each horse (13). The Akhal-Teke is currently listed as Threatened by the American Breed and Livestock Conservancy (ABLC) and Vulnerable by the Equine Survival Trust (EST) (14, 15). Threatened constitutes "fewer than 1,000 annual registrations in the United States and estimated global population of less than 5,000" by the ABLC while Vulnerable, according to the EST, means that the breed has between 500-1,500 active adult breeding mares globally (14, 15). 6 The Akhal-Teke breed was first introduced to the United States in 1979 by Phil and Margot Chase. This husband and wife team bought two horses, the stallion 1276 Senetir and the mare 2875 Oliva, at an auction in Moscow and imported them to Virginia. They also founded the Akhal-Teke Association of America (ATAA) in 1982 in the midst of the success of their breeding program (16) . Both purebreds and crossbreds were registered with the ATAA (13). As of 1997, 1276 Senetir had sired 61 foals, 32 of which were trained for competition, three of those receiving the top grade available in eventing and one attaining the top level in dressage (16). In 1995, the International Association of Akhal-Teke Horse Breeding (MAAK) was established and is currently headed by Saparmurad Niyazov, the president of Turkmenistan as well (13) . This organization is in charge of all advice on breeding for the Akhal-Teke. Inbreeding The number one enemy of genetic diversity in a population is inbreeding. Inbreeding is the mating of two related individuals. Relation is assessed by common ancestry, for example, sharing a great-grandparent (17) . The closer the relation is to the current generation, the more detrimental the pairing. For instance, while neither of these pairings is optimal by any means, mating a mother to her son is far worse than mating the son's great-grandmother to him. The mother is only one generation above her son while the great-grandmother is three generations away. Inbreeding is actually an inconvenient truth on some level for all populations (17). Frequently the number of nonrelated mates is insufficient to choose from in order to ensure the smallest amount of damage; therefore the most distantly related matings are the most desirable. A small amount of naturally occurring inbreeding is known to occur in populations. The problem arises when there is continued inbreeding resulting in an excess of homozygotes in the population, meaning there will be a large 7 proportion of individuals with identical copies of the same allele for a given gene from a common ancestor (18) . Reasons for inbreeding vary but in the horse community the most universal reason is to achieve a "perfect" specimen based on the recorded standards of the breed organization. Breeders use a method called linebreeding where they seek to increase one individual's genetic contribution to the population (19). This individual is seen as superior to others and has the qualities and traits desired to be passed on to future generations. "Popular Sire Syndrome" is a result of this type of breeding, as those sires who are physically attractive and accomplished are the most requested breeding partners. Therefore, they contribute disproportionately to the gene pool causing a genetic bottleneck (20). This thinking is a hard habit to break as the idea of the "perfect horse" is ever changing and always just out of reach. Breeders often fear they will not find such a great sample Akhal-Teke again and attempt to use the horse as much as possible while they have it (19). Also, the idea of being able to "double up" on good genes to create the best animal is a common yet flawed logic system employed by many (17). Many breeders are hesitant to equate linebreeding to inbreeding because of the negative connotation given to the latter however they are the same thing and have the same consequences. A serious result of this practice is known as inbreeding depression or the reduction in the population's fitness by the accumulation of recessive deleterious traits. Conformation issues, sterility, reduced lifespan and specific disease susceptibility are a few of the consequences (21). Some diseases found in Akhal-Tekes include Wobbler Syndrome, a cervical spine malformation, Hairless Foal Syndrome, a lethal recessive condition where hairless foals are born and die shortly after, and Degenerative Suspensory Ligament Desmitis (DSLD), a degenerative connective tissue disorder (22). To avoid these issues and help maintain the vigor of the breed the encouraged 8 practice should be that of variety. Strictly adhering to this philosophy should help to restore genetic diversity. Genes are comprised of alleles and any given individual possesses two alleles per gene. These allele combinations can be classified in one of two ways: autozygous or identical by descent (IBD), or allozygous or not identical by descent (nIBD). It is important to note the concept of IBD refers to the origin of the allele rather than its chemical makeup (18). These two distinctions are helpful when examining how inbreeding is recorded. Individuals in a population are assigned a number, which is calculated based upon their parentage, called an inbreeding coefficient (F). This is "the probability that the two alleles of a gene in an individual are [autozygous] (18)." Within a population, a non-inbred individual would have an F value of zero and an inbred individual would have an F value of greater than zero depending on the number of common ancestors. Determining if an allele is IBD requires knowledge of both sides of the family. An allele is IBD only if inherited from both parents who have ancestors in common. There are three specific steps used when calculating F from a pedigree. The first step requires locating all common ancestors. Step two calls for tracing all possible pathways for gametes to an individual from the oldest common ancestor(s) through each of the two parents (18). Each path is mutually exclusive. The third step involves the calculation of "the probability of autozygosity in [the individual of interest] due to each of the paths in turn (18)." The formula is as follows : Where F equals the inbreeding coefficient and is the sum of the paths through which an individual can derive identical alleles from the ancestors and n= the number of individuals in the paths. The 1+FA is correction factor for the inbreeding coefficient of the common ancestor in the path (23). 9 CHAPTER 2 Materials and Methods Materials Pedigree data for the calculation of inbreeding coefficients and the generation of the pedigrees was obtained from articles written by Milena Stoszek (one of the primary USA registry contacts for MAAK) posted by Akhal -Teke UK (www.akhaltekeuk.com) and Akhal-Teke Canada (www.akhal-teke.ca) and her own website Akhal-Teke Ranch (www.akhaltekeranch .com) (24-26) . Additional pedigree data was compiled from : KaraKum Akhal-Tekes (www.akhalteke.net). The Turanian Horse Website (www.turanianhorse.org), Stavropol Stud (www.stavropol-teke.com), and Akhal-Teke Est (www.akhalteke.ee) (27-30). Line founders were determined using the information provided by the Akhal-Teke Association of America' s pedigree (http://akhal-teke.org/pedigree.html) (31) . Methods Data was assembled in two databases, MyHeritage Family Tree Builder Version 4.0 (www.myheritage.com) and GenoPro 2007 (www.genopro.com) (32, 33). The data set was placed in a Microsoft Excel 2007 spread sheet including name, year and location of birth, sire, dam, color and studbook number. The data was reformatted for importing into MyHeritage Family Tree Builder. This program converts the .xlsx file format to the .gno file type readable by GenoPro. The new file was then opened in GenoPro which generated pedigrees. Additional data including founders and other progeny was added directly to MyHeritage Family Tree Builder and GenoPro 2007. 11 CHAPTER 3 Results The entire data set consisted of 527 individuals, 280 males and 247 females, shown in Appendix B. GenoPro generated one large pedigree involving 440 individuals assembled into 235 families and 25 smaller pedigrees encompassing the remaining 87 individuals in 31 families, see Table 1 and Appendices C and D (33). A family is defined as a mating and its resulting progeny. The pedigrees generated revealed more unrelated males and females in the group of smaller pedigrees than in the large pedigree, see Table 1. Sixteen diagrams were drawn for chosen line sires. The sire lines 26, 448, 432, 569, 697, 828, 721 and 914 all had progenitor sires, those stallions at the base of a split in the line, seen in Figure 1. The sire lines 269 and 779 were considered their own progenitor stallions in this examination as they themselves were at the base of the split. The presence of loops indicated inbreeding and those loops were used in the creation of the F formulas seen in Table 2. Six of these diagrams show no loops but were straight stallion line descent (Figure 1). Eight of the sixteen had a single loop and two sire line diagrams (779 and 914, a progeny of 779) revealed two loops or a second path increasing inbreeding for subsequent generations (Figure 1). The smallest of these belongs to 2a Boinou with 4 individuals resulting in an inbreeding coefficient (F) of zero (Figure 1, Table 2). The largest diagram was 914 Polotli with 24 horses, two loops, and an F of 0.0000458 (Figure 1, Table 2) . The mean number of individuals was 10.94 per sire line analyzed and the mean number of generations was 8.08. 12 Those diagrams without loops were assigned an F value of zero because they lacked any visible inbreeding. The F values for diagrams with loops were calculated using the formula above in Chapters 1 and 2. The number of individuals, excluding the Progenitor Sire at the base of the loop, directly involved with a given loop becomes the exponent (x) used in the F* formula (23). These F* values were then used to calculate the Sire Line Fvalues of (~rF* as shown in Table 2. Representative horses were chosen from each sire line for further inbreeding analysis. These individuals were the result of direct sire lines with unrelated female matings. The F values in Table 3 reveal the positive effects of breeding to unrelated individuals. Note these examples did not include additional loops in their lines. 13 Figure la. Diagrams representing line sires' ancestry for individuals 2a, 44, 165,9, 213, 100,432, 697,828,26, 569, and 269. Each dot represents a male individual unless otherwise noted with a 9 (female). When read from top to bottom, each line represents a parent-child relationship. 1\ refers to the line sire whose lineage is being examined. 151 i q F~O ku+ly So.kar 4-32F=O.OISb t:~Sa)::o.. ( ol~ Fc:: 0.000122. 2h F::O.03/3 14 Figure lb. Diagrams representing line sires' ancestry for individuals 448, 721, 779, and 914. Each dot represents a male individual unless otherwise noted with a 9 (female). When read from top to bottom, each line represents a parent-child relationship . being examined . 12.1 F =o.CX)'~,c1/ 1\ refers to the line sire whose lineage is 15 Table 1. Pedigree data including family information from GenoPro 2007 and information pertaining to related and unrelated individuals. " ..$' ~ § ',..~ .~ ..$' 1 '4 -& oS ~ ~ ~ ~ f ~ ~ "b ~ ~ ~ ~ ~ ! .; ~ ~ " ~ • .:.> Pedigree Large lJ § ~ § ~ ~ ~ f ~ G ~ ~ .; ~ 'a7 4. 'a7 ~ § o~ tI" ~ .:::> ~ o~ 440 235 224 216 16 7.100A, 179 82.900A, 87 31 56 31 25 29% 31 1000A, Individuals Table 2. Inbreeding calculations (F) for predominant sire line horses based on F* values for recognized ancestral/progenitor sires. Sire Line Sire Line Horse Steps from Progenitor Siret Sire Line Fvalue Progenitor Siret Progenitor Siret F* Formula F*Value Boinou Bek-Nazar-Dor Mele-Kush Ak-Belek Sap-ar-Khan EI n/a n/a n/a n/a 0 0 0 0 0 0 0.0156 0.000122 0.000122 0.0313 0.0156 n/a n/a n/a n/a n/a n/a n/a n/a (1/2)3(1+FBA) (1/2)12(1 +FI($) (1 2)12(1 +FJ(5) (1/2)4(1 +Fa) (1/2)4(1+FB) n/a n/a 0.0625 0.0625 (1/2)4(1+F a) 0.0625 # 2a 44 175 9 213 nfa 697 828 26 569 Gelishikli Fakirp,elvan Arab Skak n/a 3 1 1 1 2 269 Everdy-Teleke n/a 0.0625 448 721 2 4 0.0156 0.00391 779 Kir-Sakar Kaplan (Gaplan) Peren 244 Top-orbai 629 Fakir-Sulu 629 Fakir-Sulu 4a Ag-Ishan 269 EverdyTeleke 269 EverdyTeleke 4a Ag-Ishan 4a Ag-Ishan n/a 0.0000916 779 Peren 914 Polotli 1 0.0000458 779 Peren 0.0625 0.0625 [(1/2)15(1 +FB))+ [ 1/2 14(I+FB)] [(1/2)15(1 +Fa)]+ [(1/2)14(1 +Fa)] 0.0000916 0.0000916 * Calculated under the assumption that 2a Boinou (B), la Baba Achun (BA), and Kutly Sakar (KS) have Fvalues approaching zero. t Most ancestral sire for which inbreeding can be formally calculated . 16 Table 3. F values for example stallions from each ofthe sire lines with an F ~ Founding Line 721 Kaplan (Gaplan) o. F of Founder Example Stallion Steps 1 0.00196 448 Kir-Sakar 0.0156 736 Keimir 1 0.0078 569 Skak 0.0156801 Spesivii 1 0.0078 26 Arab 0.0313 677 Alam 2 0.00783 432 Karl avach 0.0156 879 Dorsekil 2 0.0039 828 Fakirpelvan 0.00391 904 Melekush F value 0.000122 1248 Orlan 2 0.0000305 779 Peren 0.0000916 1276 Senetir 2 0.0000229 914 Polotli 0.0000458 1084Padishakh 9 2 0.0000115 269 Everdy-Teleke 697 Gelishikli 0.0625687 Arsenal 0.000122 1240 Melechan 3 0.00781 3 0.0000153 17 CHAPTER 4 Discussion and Conclusions Discussion The data set of 527 individuals covered approximately 14 generations when referring to the 2a Boinou line. The majority of males in the large pedigree are associated with this line. The large pedigree has 235 families that are all connected in one way or another through both male and female lines (Appendix O) . There were numerous occurrences of linebreeding due to the same sires being used over and over. In fact, as shown in Appendix 0, it is rare that mares are used more than once. This consistently high volume of new females may have contributed to ineffective record keeping but on the other hand, the abundance of females dramatically helped the inbreeding situation of the Akhal-Teke. The females appear to be the major source of new genetic material, see Table 1. If females were used multiple times, or as often as males, then the problem would have been compounded. Unrelated males in the large pedigree were found to be few and far between. They appeared to be randomly placed though this could mean simply they have yet to be further connected to other participants of the large pedigree or the 25 smaller pedigrees (Appendices C and O). Similarly, the 25 individual pedigrees in Appendix C are unable to be connected to the large pedigree though it is suspected that they are all involved somehow. Further investigation into these connections will allow for the addition of the smaller families. The lack of depth, usually beyond the fourth generation, exhibited by the sources available through this non-Mother Russia studbook approach contributed to this problem (26-31). When the individual pedigrees are successfully integrated into the large pedigree the inbreeding coefficients will need to be checked and possibly 18 recalculated as the addition of more horses could create new or exacerbate already existing inbreeding loops, see Figure 1 (23). Sixteen diagrams were drawn for chosen line sires. The sire lines 26, 448, 432, 569, 697, 828, 721 and 914 all had progenitor sires, those stallions at the base of a split in the line, seen in Figure 1. The sire lines 269 and 779 were considered their own progenitor stallions in this examination as they themselves were at the base of the split. The presence of loops indicated inbreeding and those loops were used in the creation of the F formulas seen in Table 2. Six of these diagrams show no loops but were straight stallion line descent (Figure 1). Eight of the sixteen had a single loop and two sire line diagrams (779 and 914, a progeny of 779) revealed two loops or a second path increasing inbreeding for subsequent generations (Figure 1). The smallest of these belongs to 2a Boinou with 4 individuals resulting in an inbreeding coefficient (F) of zero (Figure 1, Table 2). The largest diagram was 914 Polotli with 24 horses, two loops, and an F of 0.0000458 (Figure 1, Table 2). The mean number of individuals was 10.94 per sire line analyzed and the mean number of generations was 8.08. Of the sixteen sire line diagrams used for inbreeding analysis, ten had loops. Loops were a clear indication of multiple paths by which identical alleles could be inherited by descent. In the early period of Akhal-Teke breeding there were plenty of recognized individuals in the breeding pool. As regulation increased and the studbooks were closed the amount of inbreeding became amplified . More horses were bred back to those that were registered and proven causing an increase in loops. As evidenced by the very small increase of less than 1% in mare registrations recorded by the VNIIK from 1994-1997, records of female Akhal-Tekes are in short supply. For example, as seen in Appendix D, 2a Boinou has 8 matings, only one of which is actually known and recorded . 19 Conclusions When moving further down from the Progenitor Sires, the problem can either get better or worse. Table 3 shows cases where the F value diminished, meaning that inbreeding decreased . Therefore if enough generations pass without any previously used or related individuals being mated into the line in question, the F value could drop back down approaching zero. There were also individuals beyond the Progenitor Sires whose inbreeding diagrams would have exhibited just as many if not more loops leading back to the common ancestors. The lack of early female record keeping interfered directly with all inbreeding calculations. Pathways could not be established without both maternal and paternal ancestral knowledge as discussed previously. This demonstrated the importance of maintaining accurate records for both male and female contributions as without these records proper inbreeding coefficients could not be determined and breeders do not have all the necessary information to make beneficial breeding choices. Inbreeding will remain a problem in the Akhal-Teke breed. Its effects will only be mitigated by the very careful selection and integration of bloodlines in the form of both stallions and mares based on genetics rather than aesthetics. 20 REFERENCES 1. Anonymous. "History & Culture : The Akhalteke Horse of Turkmenistan" Embassy of Turkmenistan . <http://www.turkmenistanembassy.org/turkmen/history/horses.html>. [Date Accessed 5/4/2010). 2. U.K. Akhal-Teke 2007-2009. From an Interview with Stoszek, Milena, Registrar of the Akhal-Teke Association of America Studbook and USA MAAK Coordinator.<http ://www.akhaltekeuk.com/ Milenalnt.html> [Date Accessed 4/18/2010). 3. Equus Survival Trust Equine Conservation List. 2008. <www.Equus-Survival-Trust.org> [Date Accessed 5/14/2010). 4. 2009 North American Akhal-Teke Stallion Directory. Shirley Schulz, e-mail: fa rmsss@belisouth.net 5. Solovyov, D. Cult of noble horse alive in Turkmenistan. The Nation 1998 February 9; Sect C; 5 (coil). <http://news.google .com/newspa pers?n id=437 &dat=19980209&id=wtQtAAAAI BAJ&sjid= _TE DA AAAIBAJ&pg=6718,2460069>. [Date Accessed 5/14/2010). 6. Klimuk, A. "Geldy Kiarizov''<http://www.akhalteke.org/web/karakum .nsf/Articles/CC330451C63DD BBC85257497007EDBED>. [Date Accessed 4/17/2010). 7. Filipov, D. A long way to go. The Boston Globe 1998 April 5.<http://www.boston.com/news/world/ asia/articies/1998/04/05/a_long_way_to_go/>. [Date Accessed 4/14/2010]. 21 8. Stewart, K. "Volya for Geldy Kyarizov" <http ://www.squidoo.com/geldy>. [Date Accessed 5/14/2010]. 9. Amnesty International. "Central Asia: Summary of Human Rights Concerns. March 2007 - March 2008" 15p. <http://www.amnesty.org/en/library/asset/EUR04/001/2008/en/5b2d8a92-071dlldd-badf-1352a91852c5/eur040012008eng.pdf>. [Date Accessed 5/20/2010]. 10. Anonymous. "Breed Standard and Conformation" <http://www.akhal-teke.org/breed.html>. [Date Accessed 4/15/2010]. 11. Anonymous. "Breeding Selection: Pre-1900 Type" <http://www.akhalteke.info/pre-1900-type-l63-en.html>. [Date Accessed 4/14/2010]. 12. Anonymous. "Breeding Selection: Post-1900 Type" <http://www.akhalteke.info/post-1900-type1-64-en .html>. [Date Accessed 4/14/2010]. 13. Anonymous. "Akhal-Teke" Horsemanship and Horse Training Tips.<http ://www.horsemanship horsetrainingtips.com/articles/178/1/Akhal-Teke>. [Date Accessed 5/15/2010] . 14. Equine Survival Trust. <http://www.equus-survival-trust.org/>. [Date Accessed 4/28/2010]. 15. American Livestock and Breed Conservancy. <http://www.albc-usa.org/cpl/akhalteke.html>. [Date Accessed 4/28/2010]. 16. Anonymous. "Akhal-Teke Under Saddle in the U.S." <http://www.akhalteke.cc/case.html>. [Date Accessed 5/10/2010]. 17. Anonymous. "Genetics : Breeding Scheme" <http://www.akhalteke.info/breeding-schemes-1-48en.html>. [Date Accessed 4/17/2010] . 18. Hartl, DL, Clark, AG. Principles of Population Genetics. 2 nd ed. 1989. 235-280p. 19. Tave, D. Inbreeding and Broodstock Management. 1999. 50p. 22 20. Anonymous. "Breeding Selection: Type as a Breeding Criterion" <http://www.akhalteke.info/type-as-a-breeding-criterion-1-62-en.html>. [Date Accessed 4/15/2010] . 21. Anonymous. "Genetics: Inbreeding Depression" <http://www.akhalteke.info/inbreedingdepression-1-51-en.html>. [Date Accessed 4/15/2010] . 22. Anonymous. "Genetic Defects and Diseases" <http://www.akhalteke.info/genetic-defects-anddiseases-1-70-en .html>. [Date Accessed 4/20/2010]. 23. Redei GP. 1998. Genetics Manual: Current Theory, Concepts, Terms. World Scientific Publishing Company. 390-392p, 530p, 928-930p. 24. Anonymous. "Horses in the UK: Akhal Teke in the UK, Purebreds registered with the Mother Stud Book" Akhal-Teke UK Ltd. < www.akhaltekeuk.com>. [Date Accessed 5/17/2010]. 25. Anonymous. "Canadian Akhal-Tekes: Stallions and Mares" Akhal-Teke Canada. < www.akhalteke.ca>. [Date Accessed 5/17/2010] . 26. Stoszek, M. "Stallions 1, Stallions 2, Contemporary Stallions, and Mares 1" Akhal-Teke Ranch. < www.akhaltekeranch .com>. [Date Accessed 5/15/2010] . 27. Keith, JL. "PhotoTeke (various stallions)" KaraKum Akhal-Tekes. <www.akhalteke.net>. [Date Accessed 5/17/2010]. 28. Riabova, T. "Turanian Horse Website: Akhal-Teke Sire Lines in Depth" The Turanian Horse Website <www.turanianhorse.org>. [Date Accessed 5/17/2010]. 29. Kl imuk, A. "Sires and Broodmares" Stavropol Stud. <www.stavropol-teke.com>. [Date Accessed 5/18/2010] . 30. Leisson, K, Puusepp, K. "Stallions and Mares" Akhal-Teke Est. <www.akhalteke.ee>. [Date Accessed 5/18/2010]. 23 31. Laux, A (2008 Board President). "The Amazing Akhal-Tekes Pedigree" Akhal-Teke Association of America . 2008 November 4. < http://akhal-teke.org/pedigree.html>. [Date Accessed 5/17/2010] . 32. MyHeritage Family Tree Builder Version 4.0. 2006-2010. <www.myheritage.com>. [Date Accessed 3/25/2010 to 5/25/2010]. 33. Morin, D. GenoPro 2007 . Canada, 1998-2010. <www.genopro.com>. [Date Accessed 3/25/2010 to 5/30/2010]. 24 APPENDICES Page Appendix A: Akhal-Teke Breed Coat Colors 25 Appendix B: Pedigree Data 26 Appendix C: Pedigree Diagram for the Smaller Families 40 Appendix D: Pedigree Diagram for the Large Family 41 25 Appendix A: Akhal-Teke Breed Coat Colors Color Nickname Description Example Horse Dominant Black Electric Black, Raven Black, "Voronaya"* Black that glitters with blues or purples Dappled, may turn complete ly white Brown with black tips to the hairs Brown that glitters with reds and golds Dark chocolate-colored brown Golden or reddish brown Tan with or without spinal stripes a few shades darker than the coat Tan with or without spinal stripes a few shades darker than the coat that glitters with golds Cream color with a bicolor mane Very light reddish cream color t Golden with chrome light golden with t chrome light to the point of almost white with blue eyes light to the point of almost white with reddish or brownish tails, knees, hocks, and legs with blue eyes Combination of dark golden dun and perlino Olive version of dark golden dun 1021 Azal Grey Mahogany Bay Bay liver Chestnut Chestnut Golden "Bulanaya"* Dark Golden Dun Dunskin Clayback/Red Dun Palomino light Palomino Cremello " lsabella"* Perlino Cream Grulla Grulla Champagne * Russian nicknames t Facial white and high leg white with belly spot or roan patch 822 Olimpic Gul AAKT62 Astrachan AAKTl48 Gora 828 Fakirpelvan AAKT29 Doblet AAKTl Senetir AAKT130 Glozenov MV Mazan 885 Kambar AAKT146 Aliger Chakan 26 Appendix B: Pedigree Data [Excel data pages follow] 27 Appendix B: Pedigree Data Gender Horse ID# 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 F M M M M M F M M M M M M M M M M M M F M M M M M M M M M M M M M M M M M M M M M 026 Vekherzen 028 Mele 044 Tillya-Kush 100 EI 1012 Sumbar 1014 Telekush 2nd 1015 Fialka 1017 Yanichar 1021 Azal 1022 Asam 1026 Alagir 10 1028 Amelit 12 1029 Anchar 1030 Arab 1031 Akhmed 1033 Beduin 1034 Beshtau 1038 Gaisan 7 1039 Galkan 104 Gezel 1045 Gektor 12 1050 Gorlik (error numbered 1051) 1052 Gubalak 5 1053 Gumdag 1056 Dasht 1058 Dost 1060 Zergyar 18 1066 Kemal 1067 Koldun 1069 Kortik 1073 Meidanli 1075 Melgun 1076 Melekush 10830mar 1084 Padishakh 9 1085 Pazyl 1087 Parakhat 1093 Perkhat 1095 Polot 1098 Saklanma 1099 Serasker 28 10# 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 Appendix B: Pedigree Data Gender Horse 1100 Taran M 1101 Turali M 1106 Algush F 1106 Khutor 17 M 1110 Yunus M 1114 Yaran M 1115 Yasman 9 M 1117 Azan M 1120 Akbelek M 1121 Akzhol M 1122 Aksum M 1123 Aky/ly M 1124 Alada M 1126 Alaman 8 M 1127 Almaz M 1128 Alsakar M 1129 Altyn 8 M 1130 Alakhrat M 1131 Alp M 1132 Amanat 23 M 1136 Arguvan 5 M 1137 Arkadash M 1138 Arslan M 1139 Askerkhan 9 M 1140 Asman M 1140 Ball F 1141 Astrachan M 1149 Bilbil F 1149 Gay 19 M 1150 Gaigysyz M 1151 Galabeg M 1154 Galkan M 1158 Gul F 1160 Garant (Akhal 2nd) 20 M 1161 Garpun M 1161 Gyarsashli F 1162 Garsak M 1166 Gayadagy M M 1166 Gayaz 1171 Gigant M 1172 Gindarkh 13 M 29 Appendix B: Pedigree Data Gender Horse 10# 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 116 117 118 119 120 121 122 123 124 M M F M M M M F M F M M M M F F M M M M F M M M M M M M M M F F F M F M M F M M M 1179 Gunesh 22 1185 Dzhasman 1185 Elek 1188 Dzhuma 1189 Dialog 1190 Doblet 1192 Dorkush 1193 Eisona 1195 Ekemen 1195 Enysh 1198 Enyshli 1202 Karaburgut 1206 Kashman 1207 Keimir 2nd 1212 Kalinka 1213 Kamina 1219 Kuwatly 1224 Madras 1225 Mazan 1226 Maidan 6 1227 Karali 1227 Makat 1228 Maksut 2 1230 Mamuk 1231 Mangyt 1232 Marakan 1233 Maruk 1240 Melechan 1241 Melok 1242 Mergen 1243 Kachakchi 1244 Kachkir 1245 Kashan 1245 Muzar 1247 Keimirchi 12470lafchan 1248 Orlan 1249 Kelte 1249 Osman 125 Kara-Kush 12500trar 30 ID# 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 Appendix B: Pedigree Data Gender Horse 1252 Palmar M 1254 Kesketli F M 1254 Panache 1262 Koimet F 1262 Perenli M 1268 Pieter M 1269 Piyada M 1274 Satrap M 1275 Sengar M 1276 Senetir M 1277 Sentor M 1280 Singapur M 1281 Kyrmyzy F 1281 Sozdnik 31 M 1283 Sugun M M 1284 Sultanbala 1291 Tyllanur M 1293 Farukh 19 M 1295 Khadji M 1300 Melepel f 1301 Yakor M 1303 Mechta F F 1305 Moroshka 13100vgan F 1316 Paska F 1318 Peidachi F F 1321 Pobeda F 1363 Tovshan 1365 Torkush F 1376 Fergezel F F 1391 Khustka 1408 Agava F 141 Keshik-Sakar M 1410 Aza F F 1416 Aishat 142 Kizyl M F 1421 Akra F 1429 Alkeik F 1431 Altyn 1455 Gerdje F 1469 Guldjakhan F 31 Appendix B: Pedigree Data Gender Horse 10# 166 167 168 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 201 202 203 204 205 206 F F F F F F F F F F M F F M F F F F F F F F F F F F F F F F F M F F F F F F F F F 1471 Gulsary 1474 Gulyali 1484 Dertli 1520 Eisona 1526 Zardiana 1539 Kaplya 1546 Karlangach 1577 Kometa 1589lebap 1594 Mantia 16 Aigir 1603 Maya (Maisa) 1609 Melekush 161 Mamed 1616 Merkenka 20 1620 Millet 1624 Mysal 16310ktava 1639 Passia 1646 Perizad (Fistashka) 1652 Sayat 1680 Totykush 1685 Fakya 1688 Fialka 1696 Khanum 1700 Cholmaya 1702 Shokhrat 1704 Yalkym 1737 Geze 1740 Grenada 1 1748 Gulshat 175 Mele-Klish 1752 Gunym 1753 Gurat 1764 Ooka 3 1769 Dortbaital 177 Elkab 1786 Kaknlls 1795 Kelkush 1797 Kepgyr 1819 Malaizia 32 Appendix B: Pedigree Data Gender Horse 10# 207 208 209 210 211 212 213 214 215 216 217 218 219 220 221 222 223 224 225 226 227 228 229 230 231 232 233 234 235 236 237 238 239 240 241 242 243 244 245 246 247 F F F F F F F F F F F F F F F F F F F F F F F F M F F F F F F F F F M F F F F F M 1820 Medeya 1842 Rovshan 1858 Takappara 2 1867 Feya 2nd 1869 Khvatka 1871 Khunta 12 1872 Citata 4 1893 Akbella 1919 Gaya 1922 Gimnastika 1949 Dovamly 1957 Dush 1964 Etgir 1987 Korsika 1991lada 2003 Melegul 2014 Mushmula 3 2037 Pugta 2040 Saga 3rd 2056 Selbiniaz 2063 Syagul 2073 Torly 2080 Formaly 2089 Abrai 209 Sakar-Keshik 2097 Azalia 2099 Aibelek 2105 Aimal 2108 Aisuy 2109 Aikhanum 2114 Akpamyk 2115 Aksoltan 2120 Alabakhar 2129 Alchak 213 Sapar-Khan 2135 Amara 4 2158 Bazilik 2163 Bossan 2181 Garda 2194 Gerda 34 220 Sluchai 33 Appendix B: Pedigree Data Gender Horse 10# 2201 Glubina F 248 F 2246 Dabarali 249 F 2250 Damdzha 250 251 252 253 254 255 256 257 258 259 260 261 262 263 264 265 266 267 268 269 270 271 272 273 274 275 276 277 278 279 280 281 282 283 284 285 286 287 288 F F F F F F M F F M F F M F M M F F F F F M F F F F F F F F F M F M F F M M 2277 Elan 2359 Malikat 2374 Mashuk 2393 Mikuna 2394 Millet 24080sanna 244 Toporbai (Bek-Nazar-Bai) 2448 Saya 2464 Ssora 247 Tugurbai 2543 Akgozel 39 2565 Almagul 26 Arab 2602 Ataka 268 Chumuch 269 Everdy-Teleke 2733 Dunja 2752 Kalina 2778 Kushka 2800 Marishka 2803 Marsiya 8 281 Yulduz 28730lesia 2875 Oliva 2883 Pampa 2886 Parfia 2890 Pelenli 2894 Perigozel 2900 Piroshka 2939 Sonata 2955 Fargona 2a Boinou 312 Mele-Zor 347 Vezir 351 Pel 377 Skala 3a Voron (Bakharden-Kara) 4 Ag-Ishan 34 Appendix B: Pedigree Data 10# Gender Horse 289 290 291 292 293 294 295 296 297 298 299 300 301 302 303 304 305 306 307 308 309 310 311 312 313 314 315 316 317 318 319 320 321 322 323 324 325 326 327 328 329 F M M M M M M M M M M F M F M F M F M M M M M M M F M M M M F M M M M M M M M M F 420 Firyuza 432 Karlavach 435 Kara-Mashal 44 Bek-Nazar-Dor 448 Kir-Sakar 454 Koshpeli 518 Moskva 569 Skak 571 Sudan 579 Synok 596 Tezir 60 Barok 600 Tetyr 627 Dor Oepel 629 Fakir-Sulu 647 Egoza 668 Absent 670 Erke-Djan 672 Akbar 675 Aksakal 677 Alam 682 Almaz 682 Bakhar 685 Angar 687 Arsenal 69 Beifid 690 Askol 697 Gelishikli 699 Gundogar 6a Dovlet-Ishan 705 Kara-Depel 711 Erkus 721 Kaplan (Gaplan) 724 Karakir (Karakunon) 736 Keimir 746 Kirdja 747 Kirsakar 779 Peren 789 Saper 795 Serdar 8 Aiden 35 Appendix B: Pedigree Data Gender Horse 10# 330 331 332 333 334 335 336 337 338 339 340 341 342 343 344 345 346 347 348 349 350 351 352 353 354 355 356 357 358 359 360 361 362 363 364 365 366 367 368 369 370 M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M F 812 Taimaz 828 Fakirpelvan 831 Farfor 834 Fed 85 Dor-Bairam 850 Azamat 854 Akbelek 2nd 858 Ametist 860AIgyr 861 Akhal 862 Bakhar 866 Vatanchi 87 Dor-Depel 870 Garem 11 (Gamburg) 871 Gindukush 874 Guneshli 875 Gurgen 876 Derbi 2nd 879 Dorsekil 882 Kalkan 883 Kaltaman 885 Kambar 889 Kemer 892 Keppan 894 Kerven 895 Kermek 2nd 896 Kerogly 9 Ak-Belek 902 Kyrgi 903 Meleke 904 Melekush 909 Ovlaksakar 2nd 910 Opal 913 Polet 914 Polotli 915 Posman 917 Sektor 918 Senagat 919 Sere 930 Finish 931 Sinitza 36 Appendix B: Pedigree Data 10# Gender Horse M 932 Khorgos 2 371 372 373 374 375 376 377 378 379 380 381 382 383 384 385 386 387 388 389 390 391 392 393 394 395 396 397 398 399 400 401 402 403 404 405 406 407 408 409 410 411 M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M F F F M M F F 933 Khorog 935 Yulduz 939 Adat 942 Alvan 943 Arslan 944 Gaukhar 4 945 Gerden 946 Gomon 4 949 Dornazarbek 950 Enysh 960 Mukhtar 961 Parasat 962 Pasyl 967 Pudok 968 Pursat 969 Rokot 970Sandjar 971 Khalif 972Shakhi 973 Abdulrakhman 974 Aigytly 975 Aktash 1 977 Arkadag 981 Garazat 983 Goklen 985 Dagestan 986 Derbent 987 Dzhagitai 989 Karader 990 Karaman 995 Kerzi (Kerrou) 997 Markiz 999 Munir Ak Kyz Ana-Kuli Arma Aryk (Arik) Ata-Khodjanok Young Ayla Bakkara 37 10# 412 413 414 415 416 417 418 419 420 421 422 423 424 425 426 427 428 429 430 431 432 433 434 435 436 437 438 439 440 441 442 443 444 445 446 447 448 449 450 451 452 Appendix B: Gender M M F M M M F F F F M M M F F F F F F F F F F M F F M F F F M F F F M F F F F F F Pedigree Data Horse Bek-Nazar-AI Enilmez Everdy-Teleke Farkhat Niderbai Father of Sailan lUas-Kara Inglis-Kurro Kepter-Tai Khotcha-Kor Komek-Cherkez-li Kurt-Bai lelianing-Chep Mantia Mother of 161 Mamed Mother of 209 Sakar-Keshik Mother of 26 Arab Mother of 281 Yulduz Mother of 2a Boinou Mother of 87 Dor-Depel Mother of Sailan Movlian-Berdy Mulla-Begench (Durdy-Niaz) NN Atbasar Oraz-Niaz-Karadashli Ovez-Mukhamed-Khan Sailan 1726 Arfa 1389 Hasma 16270vadan 726 Karat 1880Yalma 1705 Yarum 1488 Djakhan 801 Spesivii 1252 Kepderi (lachin) 1548 Kaska 9 1132 Asmanall 1636 Palma 604 Djeren 801 Mama Gezel 38 Appendix B: Pedigree Data Gender Horse 10# 453 454 455 456 457 458 459 460 461 462 463 464 465 466 467 468 469 470 471 472 473 474 475 476 477 478 479 480 481 482 483 484 485 486 487 488 489 490 491 492 493 F F F F F F F F F F M F M F M F M F M F M F M F F M F F F F F F M F M F F F M F M 1108 Alkeik 1417 Akacia 1693 Fortuna 1388 Frakcia 1165 Depel 1375 Fauna 1434 Aragva 1760 Djakhan Mother of Everdy-Teleke Mother of Nasar Belek Dor Nasar Belek Dor Mother of Bairam Kor (Bairam Corinthians) Bairam Kor (Bairam Corinthians) Djaj Bai Khait Bai (Schaschibai Chaitbaj) 5a Dschaschi Bai (Djaji Bai) Mother of Keppan Keppan Mother of 8a Tschep Mele (Mele Chep) 8a Tschep Mele (Mele Chep) Mother of Enilmez Mele Khadjdi Nur (Mele Hadzhi Nur) Shikhli Nur Kaza 1a Baba Achun Mother of Mele Khadjdi Nur (Mele Hadzhi Nur) Mother of Kurban Nijas Kurban Nijas Hodzda Boll Mother of 6a Dovlet-Ishan Ishan Sha Durdi Aj Dschamal 124 Gul Dzhahan 133 Kara Enym Mother of 133 Kara Enym Ak~Yal Mele Mother of Bek-Nazar-AI Mother of Ak-Yal Mele Mother of 420 Firyuza Kaib Bai Mother of Komek-Cherkez-li Sultan Guli 39 10# 494 495 496 497 498 499 500 501 502 503 504 505 506 507 508 509 510 511 512 513 514 515 516 517 518 519 520 521 522 523 524 525 526 527 529 530 531 532 533 534 Appendix B: Pedigree Data Gender Horse F M F M F F F M F F F F F F F M M F M F M F F M F F F M M F M F F F F F F F F F Polot Karamtscha Mother of lelianing-Chep Kutly Sakar Mother of Karamtscha 1388 Fraction 1325 Sadap 996 Mag 1665 Sulgun 1292 Makhim 1381 Filfil Silma 1378 Festival 1689 Fialka 1359 Tayar (Tkachikha) 741 Kerem 338 Birgut Mother of 338 Birgut Ata-Khodjanok Old Mother of Ata-Khodjanok Young Fortingbrass - TB Mother of Ata-Khodjanok Old Mother of 198 Posman 198 Posman Mother of Khotcha-Kor Mother of 687 Arsenal Mother of Shikhil Nur Kaza 172 Mele Ker Suncheli Mother of 268 Chumuch Suncheli Mother of Suncheli 831 Mah Atla-Bally Mother of 677 Alam Mother of Khustka Khustka Mother of 801 Spesivii 1355 Talent 1524 Enish 40 Appendix C: Pedigree Diagram for the Smaller Families o o 0 1795 KeI<ush 0 1026 2106 r 00 0 0 0 0 0 0 1872 1073 1740 1053 2105 1058 2448 1052 1869 Aisuy ~-..... 1fT ,,1 r r , ~ ~y~ To Y YYV T TPo Y TT ToTo "';'" 1129 1100 Tar.., Tyllru 8 o o 0 MoIher of 690 Ask'" SaIII'l o 0 0 0 0 0 675 Aksakal o - 1242 (Akhal 2nd) 20 Alyn 1291 "t" 0 0 _ar Khaqi Mergen o o o 1095 2109 1126 1140 Asmari o - 0 0 1281 AIsnan 0 1110 2374 8 o o o o 1224 o 2900 1268 Pieter 1117 Azan 1124 -'- 1188 DzIuna 209 1262 Perri Sakar- K_ o 0 B50 0 0 973 1589 2056 y"y 942 1700 1120 Akbelek 1138 2114 1130 AlaklTat