649:
237:
470:
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67:
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488:, causing the coat color to be diluted from red to gold, usually without primitive markings. Visually, a bay dun is a tan-gold color, somewhat darker and less vivid than the more cream or gold buckskin, and duns always possess primitive markings. Today, pedigree analysis, DNA testing, studying possible offspring, and the vividness of primitive markings are used to determine whether a horse is a dun.
229:
1427:
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50:
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is still functional, and is still produced in most cells, but not expressed in the hair cortex. Where the coat is diluted, the color is not uniform throughout each hair, but rather is more intense on the outward-facing side of the hair shaft and lighter underneath. In the darker areas, where the
103:
the shade of the undiluted base coat color. A dun horse always has a dark dorsal stripe down the middle of its back, usually has a darker face and legs, and may have transverse striping across the shoulders or horizontal striping on the back of the forelegs. Body color depends on the underlying
509:
To further confuse matters, it is possible for a horse to carry both dun and cream dilution genes; such horses with golden buckskin coloring and a complete set of primitive markings are referred to as a "buckskin dun" or a "dunskin". On such horses, the light-shaded primitive markings are most
752:
expression mirrored the pattern of pigment deposition in the hair, that is, TBX3 was found wherever the pigment was not. TBX3 was not found in the hair cortex keratinocytes from non-dun horses nor in those from the dorsal stripe of dun horses. However, all of the horses had a thin
484:, in that both colors feature a light-colored coat with a dark mane and tail. In particular, buckskins with non-dun 1 primitive markings can easily be confused with dun. Genetically, a bay dun is a bay horse with the dun gene. A buckskin is bay horse with the addition of the
703:. It is thought that the non-dun2 genetic mutation (as well as the development of chestnut base color) occurred after domestication. Ancient DNA from a horse that lived about 43,000 years ago, long before horses were domesticated, carried both dun and non-dun1 genes.
635:
primitive markings occur, the hair shaft is of uniform color. One of the researchers involved in the study said it could be called a "microscopic spotting pattern". This phenomenon is new to science and has not been observed in rodents, primates, or carnivores.
610:
The dun dilution effect is caused by pigment only being placed in a part of each hair. Specifically, hairs from diluted areas only have pigment along one side of them, while hairs from darker parts such as the dorsal stripe have pigment all the way around.
626:. When functional, it creates dun coloring, including the primitive markings, and when recessive, a horse is not dun. In humans and lab mice, TBX3 is critical to development. Abnormalities are linked to a collection of developmental defects called
557:
gene; however, at least one study found a statistically significant variation in the shade of dilution depending on whether one or two copies of the dun gene are present. Two non-dun parents cannot produce a dun foal. Horses that are non-dun1
340:, is a smoky, blue-gray to mouse-brown color and can vary from light to dark. They consistently have black points and they often have a dark or black head. The primitive markings are usually all black. Genetically, the horse has an underlying
495:, which is genetically a bay horse with two copies of the cream gene, which creates a horse with a cream-colored body but a reddish mane and tail. However, perlinos usually are significantly lighter than red dun and have blue eyes.
399:
does not significantly lighten black hair, though it may have a subtle effect, and thus a single copy generally has no visible effect on a grullo, either. Conversely, double copies of the cream gene create very light-colored horses
116:
base coloration are a smoky gray. Manes, tails, primitive markings, and other dark areas are usually the shade of the undiluted base coat color. The dun gene may interact with all other coat color alleles.
183:
can also be considered a variant of dun where the dilution is so extreme it turns the hair nearly white, and the primitive markings (like the striped leg barring) extend across the entire body.
506:, which also intermingles light and dark hairs, the color does not change to a lighter shade as the horse ages. With a dun, the hair color is one solid shade and remains so for life.
773:, were found only in the pigmented areas of the hair. This indicates that the hair follicles of dun and non-dun horses have different distributions of pigment-producing cells.
648:
684:
There are two forms of non-dun color, non-dun1 and non-dun2, caused by different mutations. Non-dun1 horses have some primitive markings, while non-dun2 horses do not.
362:
Facial mask, a darker area around the nasal bone and forehead, sometimes making the head close to the undiluted color, some patterns colloquially called "Cobwebbing."
517:
Countershading such as light dorsal stripes resulting from the presence of the gene nd1 (see section below) may be difficult to detect on light-colored horses.
427:. Non-dun1 horses have no dun color dilution but may keep primitive markings, while non-dun2 horses have neither the dun color dilution nor primitive markings.
1293:
Cieslak, Jakub; Brooks, Samantha Ann; Wodas, Lukasz; Mantaj, Weronika; Borowska, Alicja; Sliwowska, Joanna Helena; Ziarniak, Kamil; MacKowski, Mariusz (2021).
1459:
473:
This is a buckskin horse with no known dun genetics, showing a countershading stripe that could be confused with dun. This may be an example of non-dun1.
434:
breed, which is predominantly dun, uses unique
Norwegian-based terminology to distinguish between the different shades of dun horses. "Brown dun", or
302:
is the most common type of dun, and has a tan or gold body with black mane, tail, and primitive markings. Genetically, the horse has an underlying
477:
Historically, before modern genetic studies distinguished between alleles, diluted colors were sometimes lumped together and simply called "dun".
872:
deletion is a more derived allele. Nucleotide diversity across the flanking regions of chromosome 8 for the various alleles indicates that the
580:
horses. The primitive markings from non-dun1 are more visible on a bay or chestnut horse; they blend in on a black. A horse with two copies of
1244:
320:
coat color, acted upon by the dun gene. Thus, as there is no black on the horse to be affected, the undiluted underlying color is red.
1397:
660:
108:. A classic "bay dun" is a gray-gold or tan, characterized by a body color ranging from sandy yellow to reddish brown. Duns with a
1452:
1431:
1295:"Genetic Background of the Polish Primitive Horse (Konik) Coat Color Variation—New Insight into Dun Dilution Phenotypic Effect"
630:, and the null allele (being unable to produce any TBX3 at all) is thought to be embryonic lethal. In non-dun horses, the TBX3
833:
316:, is a light tan coat with reddish instead of black points and primitive markings. Genetically, the horse has an underlying
785:
were found all the way around the hair in non-dun horses, but only on the pigmented side in dun horses. The region where
514:
that also carries dun, showing primitive dorsal striping or leg bars indicative of a red dun may be called a "dunalino."
416:
also carries the dun gene, it also will be cream-colored, with primitive markings not visible to any significant degree.
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may have some asymmetry in pigment distribution, producing primitive markings, but to a lesser degree than dun horses.
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781:, a molecule required for melanocyte migration and survival in the skin and hair follicle. Keratinocytes expressing
627:
236:
1988:
1969:
1938:
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1903:
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219:
105:
1026:"Regulatory mutations in TBX3 disrupt asymmetric hair pigmentation that underlies Dun camouflage color in horses"
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at chromosome 8 base pair 18,226,905, which appears to be sufficient to cause non-dun1 coloration. In addition,
373:, less common in horses, and if present, often faint, usually only visible on a short summer coat, if at all.
1958:
419:
Dorsal striping alone does not guarantee the horse carries the dun gene. There two types of non-dun, called
1877:
469:
109:
1993:
1710:
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279:(red dun). It is more difficult to recognize when combined with other dilution genes or if affected by
148:
1135:
Adalsteinsson S. (May 1978). "Inheritance of yellow dun and blue dun in the
Icelandic toelter horse".
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was significantly downregulated in non-dun horses compared to dun horses, while the neighboring gene,
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Imsland F, McGowan K, Rubin CJ, Henegar C, Sundström E, Berglund J, et al. (February 2016).
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Frosting: light hairs found on either side of the mane and on both sides of the dock of the tail.
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Horizontal striping on the back of forelegs, common on most duns, although at times, rather faint
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Veterinary
Genetics Lab, University of California, Davis. Web page accessed December 4, 2009
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261:
160:
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927:"Genotypes of predomestic horses match phenotypes painted in Paleolithic works of cave art"
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Dorsal stripe down the center of the back, along the spine, seen almost universally on duns
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Genetic analysis and DNA sequencing results published in 2015 link dun color to the
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1336:"Mouse TBX3 mutants suggest novel molecular mechanisms for Ulnar-mammary syndrome"
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was not expressed was similar to, but not exactly the same as, the region where
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has a 1,609 bp deletion and another very near 8 bp deletion. Comparison with
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The location of TBX3 expression may also determine the striping pattern of
545:) horses do not have dun dilution but may exhibit some primitive markings.
1426:
1068:"A horse of a different color: Genetics of camouflage and the dun pattern"
1953:
1817:
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1565:
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Monogenic Traits
Associated with Structural Variants in Chicken and Horse
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451:
401:
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Frank DU, Emechebe U, Thomas KR, Moon AM (2013-07-02). Dettman R (ed.).
746:, was expressed in about the same amount. In dun horses, the pattern of
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is only visible on bay and chestnut coats, and then to a lesser degree.
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at chr. 8: 18,227,267. However, that SNP was also found in some dun
736:, which are both known to be involved in hair follicle development.
728:
is predicted to include binding sites for the transcription factors
498:
Grullos are sometimes confused with roans or grays. However, unlike
197:
49:
510:
noticeable during the summer months, when the winter hair sheds. A
502:, dun has no intermingled black and white hairs, and unlike a true
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effect, lightening the body coat, but has less of an effect on the
1933:
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Bay + dun + cream gene (single copy) = "dunskin" or "buckskin dun"
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553:) horses have neither dilution nor primitive markings. Dun is a
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are found in a region of equine chromosome 8 whose only gene is
770:
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Other variations result from the interplay of additional genes:
35:
1441:
876:
mutation most likely occurred on a chromosome that already had
480:
The dun gene, when on a "bay dun" horse, can closely resemble
680:
Bay horse with visible dorsal stripe, an example of non-dun1.
706:
The non-dun mutations appear to "disrupt the function of a
446:- literally "gray" - is a grullo, buckskin duns are called
95:. The dun gene lightens most of the body while leaving the
907:. Veterinary Genetics Lab, University of California, Davis
365:
Transverse stripe, a crosswise stripe along the shoulders
141:
are dun, and several closely related species in the genus
458:
A cremello, perlino or smoky cream is called "white" or
1398:"Genetics of camouflage and the Dun pattern in horses"
793:
was expressed. TBX3 is not thought to directly affect
573:
horses typically have clearer primitive markings than
591:
in that it acts on any coat color. In contrast, the
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1269:"A Horse Of A Different Color: Buckskins And Duns"
654:Dorsal stripe and light guard hairs on a dun horse
112:base may appear a light tan shade, and those with
41:A bay dun, also called a "classic" or "zebra" dun
1225:
1223:
1221:
840:that is most common in domestic horses, where a
87:that affects both red and black pigments in the
699:modifiers of the base colors bay and black for
267:Dun visibly affects all the three base colors,
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8:
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412:). Thus, if a horse with two cream dilution
388:(single copy) = "dunalino" or "palomino dun"
147:show dun characteristics. These include the
724:during hair growth." The region deleted in
348:Another characteristic of the dun gene are
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194:mutations were found in any other equids.
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599:is an incomplete dominant which must be
595:acts only on black-based coats, and the
1182:"More about Dun and Primitive Markings"
889:
644:
344:coat color, acted upon by the dun gene.
306:coat color, acted upon by the dun gene.
125:Dun is believed to be the ancestral or
491:A red dun may also be confused with a
450:or white dun, and a "dunalino" (dun +
1149:10.1093/oxfordjournals.jhered.a108913
1110:"Introduction to Coat Color Genetics"
925:Pruvost M, et al. (2011-11-07).
761:was expressed. Two markers of mature
587:Dun has a stronger effect than other
369:to the dorsal stripe, very common in
7:
352:. Dun traits include the following:
264:of the mane, tail, ears, and legs.
1396:Waara, Anneli (December 21, 2015).
1074:(Press release). December 21, 2015.
25:
868:in other species showed that the
1516:
1425:
1206:. Norwegian Fjord Horse Registry
659:
647:
603:to be fully expressed, and when
65:
48:
34:
1239:(Thesis). Uppsala. p. 42.
860:, so is not necessary for dun.
397:cream gene on a black base coat
171:, and an extinct subspecies of
1402:SciLifeLab, Uppsala University
834:single nucleotide polymorphism
454:) is called a "yellow dun" or
271:(bay, classic, or zebra dun),
1:
695:are thought to have been the
1361:10.1371/journal.pone.0067841
905:"Dun Dilution - Direct Test"
836:compared to the version of
584:lacks primitive markings.
275:(mouse dun or grullo), and
163:, an extinct subspecies of
2010:
1970:Category:Horse coat colors
1939:Endothelin receptor type B
1904:Equine coat color genetics
989:Smithsonian's National Zoo
689:domestication of the horse
666:Transverse shoulder stripe
232:Dorsal stripe on a red dun
220:Equine coat color genetics
217:
1967:
1696:
1586:
1555:
1514:
691:, dun, non-dun1, and the
205:: Dun is thought to be a
133:appearing in prehistoric
1230:Imsland, Freyja (2015).
1170:Cream - Horse Coat Color
717:in a specific subset of
708:transcriptional enhancer
1959:Melanocortin 1 receptor
1204:"Color in Fjord Horses"
952:10.1073/pnas.1108982108
1312:10.1093/jhered/esab034
858:Estonian native horses
681:
628:ulnar–mammary syndrome
525:There are three known
474:
249:
241:
233:
210:
129:color of horses. Many
121:Taxonomic distribution
1897:Genetics and breeding
1738:Base color variations
1667:Lethal white syndrome
1432:Dun gene of the horse
1202:Sponenberg, Phillip.
679:
472:
395:A single copy of the
247:
239:
231:
200:
1434:at Wikimedia Commons
1184:. Etalon Diagnostics
1092:"Dun Zygosity Test."
985:"Przewalski's horse"
624:transcription factor
1352:2013PLoSO...867841F
1299:Journal of Heredity
943:2011PNAS..10818626P
252:The dun gene has a
106:coat color genetics
1868:Primitive markings
1469:Equine coat colors
757:of the hair where
682:
593:silver dapple gene
475:
350:primitive markings
283:. Shades include:
258:primitive markings
250:
242:
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224:Primitive markings
211:
149:Przewalski's horse
101:primitive markings
99:, tail, legs, and
1989:Horse coat colors
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1612:
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1588:double dilution:
1557:single dilution:
1430:Media related to
1246:978-91-554-9295-3
848:is replaced with
571:non-dun1/non-dun1
529:of the dun gene:
384:Chestnut + dun +
201:Cave painting at
55:A “blue” dun, or
16:(Redirected from
2001:
1863:Point coloration
1806:Roaning patterns
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1275:. 15 August 2017
1273:Cowgirl Magazine
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937:(46): 18626–30.
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161:African wild ass
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1030:Nature Genetics
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991:. 25 April 2016
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1420:External links
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1838:Bend-Or spots
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1641:Silver dapple
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1346:(7): e67841.
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723:
722:keratinocytes
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367:perpendicular
364:
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115:
111:
107:
102:
98:
94:
90:
86:
85:dilution gene
82:
68:
58:
51:
37:
27:Dilution gene
19:
1994:Mammal genes
1954:STX17 (Gray)
1914:Horse genome
1796:Knabstrupper
1789:
1786:Varnish roan
1680:and patterns
1662:Sabino-white
1627:
1626:Blue dun or
1620:
1405:. Retrieved
1401:
1343:
1339:
1329:
1302:
1298:
1288:
1277:. Retrieved
1272:
1263:
1232:
1208:. Retrieved
1197:
1186:. Retrieved
1176:
1165:
1143:(3): 146–8.
1140:
1136:
1114:. Retrieved
1072:ScienceDaily
1071:
1036:(2): 152–8.
1033:
1029:
993:. Retrieved
988:
979:
934:
930:
920:
909:. Retrieved
877:
873:
869:
865:
861:
853:
845:
837:
832:has another
829:
821:
813:
809:
805:
801:
799:
797:expression.
794:
790:
786:
782:
774:
758:
747:
741:
737:
725:
705:
686:
683:
637:
613:
609:
605:heterozygous
586:
581:
577:
575:heterozygous
570:
563:
559:
550:
546:
542:
538:
534:
530:
524:
516:
508:
497:
490:
479:
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459:
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447:
443:
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429:
424:
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418:
394:
379:
347:
337:
333:
329:
323:
313:
309:
299:
295:
291:
287:
266:
251:
240:Leg striping
214:Color traits
191:
187:
186:Neither the
185:
165:plains zebra
143:
139:Chauvet Cave
124:
80:
78:
1909:Color breed
1764:Tricoloured
1741:(primarily
1601:Smoky cream
1578:Smoky black
1210:January 20,
763:melanocytes
755:outer layer
701:wild horses
432:Fjord horse
410:smoky cream
296:classic dun
260:and on the
248:Facial mask
137:such as in
1983:Categories
1743:UK English
1624:(see also
1571:Isabelline
1507:Seal brown
1279:2019-04-28
1188:2019-04-28
1116:2019-04-27
1112:. UC Davis
995:2019-04-26
911:2019-04-27
884:References
779:KIT ligand
715:expression
711:regulating
601:homozygous
597:cream gene
568:Homozygous
486:cream gene
465:Dun mimics
436:brunnblakk
386:cream gene
218:See also:
209:coloration
89:coat color
1888:Tiger eye
1792:Appaloosa
1790:See also
1616:Champagne
1505:included
1255:1651-6206
719:hair bulb
697:wild type
687:Prior to
500:blue roan
456:gulblakk.
338:mouse dun
300:zebra dun
207:wild type
127:wild type
72:A red dun
1934:KIT gene
1919:Wildtype
1883:Mushroom
1818:Rabicano
1759:Skewbald
1591:Cremello
1566:Palomino
1561:Buckskin
1490:Chestnut
1407:June 26,
1380:23844108
1340:PLOS ONE
1321:34432873
1060:26691985
971:22065780
878:non-dun1
874:non-dun2
870:non-dun2
862:Non-dun2
854:non-dun1
830:non-dun1
814:Non-dun1
806:non-dun2
802:non-dun1
777:encodes
726:non-dun2
582:non-dun2
555:dominant
547:Non-dun2
539:Non-dun1
521:Genetics
512:palomino
482:buckskin
452:palomino
448:ulsblakk
440:rødblakk
425:non-dun2
421:non-dun1
402:cremello
334:blue dun
318:chestnut
314:claybank
277:Chestnut
254:dilution
192:non-dun2
190:nor the
188:non-dun1
110:chestnut
81:dun gene
1924:Melanin
1858:Pangaré
1848:Cropout
1843:Brindle
1754:Piebald
1723:Tobiano
1596:Perlino
1371:3699485
1348:Bibcode
1137:J Hered
1051:4731265
962:3219153
939:Bibcode
850:thymine
842:guanine
826:adenine
824:has an
818:guanine
632:protein
527:alleles
493:perlino
414:alleles
406:perlino
371:donkeys
310:Red dun
292:bay dun
203:Lascaux
131:equines
18:Red dun
1929:Agouti
1878:Flaxen
1728:Tovero
1716:Sabino
1628:Grullo
1495:Sorrel
1378:
1368:
1319:
1253:
1243:
1157:731005
1155:
1058:
1048:
969:
959:
820:where
816:has a
640:zebras
408:, and
330:grulla
325:Grullo
181:Zebras
177:tarpan
175:, the
169:quagga
167:, the
153:onager
57:grullo
1873:Sooty
1853:Liver
1831:Other
1711:Frame
1701:Overo
1650:White
1636:Pearl
1548:Cream
1485:Black
1237:(PDF)
800:Both
795:KITLG
787:KITLG
783:KITLG
775:KITLG
713:TBX3
616:T-box
578:d1/d2
564:d1/d2
560:d1/d1
460:kvit.
342:black
298:, or
273:black
173:horse
157:kiang
144:Equus
114:black
93:horse
91:of a
83:is a
1949:PAX3
1944:MITF
1813:Roan
1794:and
1531:Gray
1526:Gray
1409:2016
1376:PMID
1317:PMID
1251:ISSN
1241:ISBN
1212:2010
1153:PMID
1056:PMID
967:PMID
931:PNAS
866:TBX3
810:TBX3
804:and
791:TBX3
771:MITF
769:and
759:TBX3
749:TBX3
743:TBX5
738:TBX3
734:MSX2
732:and
730:ALX4
620:TBX3
504:gray
430:The
423:and
281:gray
222:and
97:mane
79:The
1621:Dun
1502:Bay
1366:PMC
1356:doi
1307:doi
1303:112
1145:doi
1046:PMC
1038:doi
957:PMC
947:doi
935:108
852:in
846:dun
844:in
838:dun
822:dun
767:KIT
618:3 (
562:or
531:dun
444:grĂĄ
336:or
328:or
304:bay
288:Dun
269:bay
1985::
1400:.
1388:^
1374:.
1364:.
1354:.
1342:.
1338:.
1315:.
1301:.
1297:.
1271:.
1249:.
1220:^
1151:.
1141:69
1139:.
1125:^
1099:^
1082:^
1070:.
1054:.
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1034:48
1032:.
1028:.
1004:^
987:.
965:.
955:.
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933:.
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892:^
880:.
812:.
765:,
642:.
622:)
551:d2
543:d1
404:,
294:,
179:.
159:,
155:,
151:,
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1461:e
1454:t
1447:v
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1309::
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1191:.
1159:.
1147::
1119:.
1062:.
1040::
998:.
973:.
949::
941::
914:.
549:(
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533:(
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