638:
226:
459:
650:
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56:
234:
218:
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1507:
477:, 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.
39:
187:
25:
623:
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
92:
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
498:
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
741:
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
473:, 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
692:. 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.
624:
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.
599:
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.
615:. 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
546:
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
329:, 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
484:, 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.
388:
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
105:
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.
172:
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.
495:, 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.
762:, 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.
673:
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.
351:
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."
506:
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.
416:. 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.
1282:
Cieslak, Jakub; Brooks, Samantha Ann; Wodas, Lukasz; Mantaj, Weronika; Borowska, Alicja; Sliwowska, Joanna Helena; Ziarniak, Kamil; MacKowski, Mariusz (2021).
462:
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.
637:
423:
breed, which is predominantly dun, uses unique
Norwegian-based terminology to distinguish between the different shades of dun horses. "Brown dun", or
291:
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
466:
Historically, before modern genetic studies distinguished between alleles, diluted colors were sometimes lumped together and simply called "dun".
861:
deletion is a more derived allele. Nucleotide diversity across the flanking regions of chromosome 8 for the various alleles indicates that the
569:
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
1448:
309:
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.
1386:
97:. 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
1420:
1284:"Genetic Background of the Polish Primitive Horse (Konik) Coat Color Variation—New Insight into Dun Dilution Phenotypic Effect"
619:, and the null allele (being unable to produce any TBX3 at all) is thought to be embryonic lethal. In non-dun horses, the TBX3
1927:
1233:
305:, is a light tan coat with reddish instead of black points and primitive markings. Genetically, the horse has an underlying
774:
were found all the way around the hair in non-dun horses, but only on the pigmented side in dun horses. The region where
503:
that also carries dun, showing primitive dorsal striping or leg bars indicative of a red dun may be called a "dunalino."
405:
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.
1589:
1584:
1579:
770:, a molecule required for melanocyte migration and survival in the skin and hair follicle. Keratinocytes expressing
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649:
1015:"Regulatory mutations in TBX3 disrupt asymmetric hair pigmentation that underlies Dun camouflage color in horses"
817:
at chromosome 8 base pair 18,226,905, which appears to be sufficient to cause non-dun1 coloration. In addition,
822:
1917:
362:, less common in horses, and if present, often faint, usually only visible on a short summer coat, if at all.
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Dorsal striping alone does not guarantee the horse carries the dun gene. There two types of non-dun, called
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1977:
1958:
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616:
268:(red dun). It is more difficult to recognize when combined with other dilution genes or if affected by
208:
94:
1124:
Adalsteinsson S. (May 1978). "Inheritance of yellow dun and blue dun in the
Icelandic toelter horse".
729:
was significantly downregulated in non-dun horses compared to dun horses, while the neighboring gene,
1220:
1947:
1056:
1013:
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.
348:
Horizontal striping on the back of forelegs, common on most duns, although at times, rather faint
1866:
225:
98:
1982:
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1655:
137:
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1083:
Veterinary
Genetics Lab, University of California, Davis. Web page accessed December 4, 2009
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927:
916:"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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1325:"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
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534:) horses do not have dun dilution but may exhibit some primitive markings.
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153:
127:
85:
1057:"A horse of a different color: Genetics of camouflage and the dun pattern"
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Monogenic Traits
Associated with Structural Variants in Chicken and Horse
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Frank DU, Emechebe U, Thomas KR, Moon AM (2013-07-02). Dettman R (ed.).
735:, 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
725:, which are both known to be involved in hair follicle development.
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is predicted to include binding sites for the transcription factors
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Grullos are sometimes confused with roans or grays. However, unlike
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noticeable during the summer months, when the winter hair sheds. A
491:, dun has no intermingled black and white hairs, and unlike a true
440:
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effect, lightening the body coat, but has less of an effect on the
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Bay + dun + cream gene (single copy) = "dunskin" or "buckskin dun"
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542:) 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
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Other variations result from the interplay of additional genes:
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mutation most likely occurred on a chromosome that already had
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The dun gene, when on a "bay dun" horse, can closely resemble
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Bay horse with visible dorsal stripe, an example of non-dun1.
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The non-dun mutations appear to "disrupt the function of a
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435:- literally "gray" - is a grullo, buckskin duns are called
186:
84:. The dun gene lightens most of the body while leaving the
38:
1012:
896:. Veterinary Genetics Lab, University of California, Davis
354:
Transverse stripe, a crosswise stripe along the shoulders
130:
are dun, and several closely related species in the genus
1281:
447:
A cremello, perlino or smoky cream is called "white" or
24:
1322:
1387:"Genetics of camouflage and the Dun pattern in horses"
782:
was expressed. TBX3 is not thought to directly affect
562:
horses typically have clearer primitive markings than
580:
in that it acts on any coat color. In contrast, the
1119:
1117:
1115:
1258:"A Horse Of A Different Color: Buckskins And Duns"
643:Dorsal stripe and light guard hairs on a dun horse
101:base may appear a light tan shade, and those with
30:A bay dun, also called a "classic" or "zebra" dun
1969:
1214:
1212:
1210:
1123:
1112:
829:that is most common in domestic horses, where a
76:that affects both red and black pigments in the
688:modifiers of the base colors bay and black for
256:Dun visibly affects all the three base colors,
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526:), produces dilution and primitive markings.
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401:). Thus, if a horse with two cream dilution
377:(single copy) = "dunalino" or "palomino dun"
136:show dun characteristics. These include the
1086:
1069:
713:during hair growth." The region deleted in
337:Another characteristic of the dun gene are
1449:
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1380:
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183:mutations were found in any other equids.
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588:is an incomplete dominant which must be
584:acts only on black-based coats, and the
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216:
185:
1375:
1218:
1171:"More about Dun and Primitive Markings"
913:
333:coat color, acted upon by the dun gene.
295:coat color, acted upon by the dun gene.
114:Dun is believed to be the ancestral or
1970:
1456:
1190:
480:A red dun may also be confused with a
439:or white dun, and a "dunalino" (dun +
1430:
1384:
1163:
1138:10.1093/oxfordjournals.jhered.a108913
1099:"Introduction to Coat Color Genetics"
914:Pruvost M, et al. (2011-11-07).
750:was expressed. Two markers of mature
576:Dun has a stronger effect than other
358:to the dorsal stripe, very common in
1535:
341:. Dun traits include the following:
253:of the mane, tail, ears, and legs.
1385:Waara, Anneli (December 21, 2015).
1063:(Press release). December 21, 2015.
13:
660:
14:
1994:
1408:
857:in other species showed that the
1505:
1414:
1195:. Norwegian Fjord Horse Registry
648:
636:
592:to be fully expressed, and when
54:
37:
23:
1316:
1275:
1250:
1228:(Thesis). Uppsala. p. 42.
849:, so is not necessary for dun.
386:cream gene on a black base coat
202:
160:, and an extinct subspecies of
1391:SciLifeLab, Uppsala University
1152:
966:
823:single nucleotide polymorphism
443:) is called a "yellow dun" or
260:(bay, classic, or zebra dun),
1:
872:
684:are thought to have been the
453:
1350:10.1371/journal.pone.0067841
894:"Dun Dilution - Direct Test"
7:
825:compared to the version of
573:lacks primitive markings.
509:
264:(mouse dun or grullo), and
152:, an extinct subspecies of
10:
1999:
1959:Category:Horse coat colors
1928:Endothelin receptor type B
1893:Equine coat color genetics
978:Smithsonian's National Zoo
678:domestication of the horse
655:Transverse shoulder stripe
221:Dorsal stripe on a red dun
209:Equine coat color genetics
206:
1956:
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680:, dun, non-dun1, and the
194:: Dun is thought to be a
122:appearing in prehistoric
1219:Imsland, Freyja (2015).
1159:Cream - Horse Coat Color
706:in a specific subset of
697:transcriptional enhancer
1948:Melanocortin 1 receptor
1193:"Color in Fjord Horses"
941:10.1073/pnas.1108982108
1301:10.1093/jhered/esab034
847:Estonian native horses
670:
617:ulnar–mammary syndrome
514:There are three known
463:
238:
230:
222:
199:
118:color of horses. Many
110:Taxonomic distribution
1886:Genetics and breeding
1727:Base color variations
1656:Lethal white syndrome
1421:Dun gene of the horse
1191:Sponenberg, Phillip.
668:
461:
384:A single copy of the
236:
228:
220:
189:
1423:at Wikimedia Commons
1173:. Etalon Diagnostics
1081:"Dun Zygosity Test."
974:"Przewalski's horse"
613:transcription factor
1341:2013PLoSO...867841F
1288:Journal of Heredity
932:2011PNAS..10818626P
241:The dun gene has a
95:coat color genetics
1857:Primitive markings
1458:Equine coat colors
746:of the hair where
671:
582:silver dapple gene
464:
339:primitive markings
272:. Shades include:
247:primitive markings
239:
231:
223:
213:Primitive markings
200:
138:Przewalski's horse
90:primitive markings
88:, tail, legs, and
1978:Horse coat colors
1965:
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1597:
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1577:double dilution:
1546:single dilution:
1419:Media related to
1235:978-91-554-9295-3
837:is replaced with
560:non-dun1/non-dun1
518:of the dun gene:
373:Chestnut + dun +
190:Cave painting at
44:A “blue” dun, or
1990:
1852:Point coloration
1795:Roaning patterns
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1264:. 15 August 2017
1262:Cowgirl Magazine
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926:(46): 18626–30.
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251:point coloration
150:African wild ass
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980:. 25 April 2016
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1409:External links
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1827:Bend-Or spots
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1630:Silver dapple
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711:keratinocytes
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356:perpendicular
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107:
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100:
96:
91:
87:
83:
79:
75:
74:dilution gene
71:
57:
47:
40:
26:
16:Dilution gene
1983:Mammal genes
1943:STX17 (Gray)
1903:Horse genome
1785:Knabstrupper
1778:
1775:Varnish roan
1669:and patterns
1651:Sabino-white
1616:
1615:Blue dun or
1609:
1394:. Retrieved
1390:
1332:
1328:
1318:
1291:
1287:
1277:
1266:. Retrieved
1261:
1252:
1221:
1197:. Retrieved
1186:
1175:. Retrieved
1165:
1154:
1132:(3): 146–8.
1129:
1125:
1103:. Retrieved
1061:ScienceDaily
1060:
1025:(2): 152–8.
1022:
1018:
982:. Retrieved
977:
968:
923:
919:
909:
898:. Retrieved
866:
862:
858:
854:
850:
842:
834:
826:
821:has another
818:
810:
802:
798:
794:
790:
788:
786:expression.
783:
779:
775:
771:
763:
747:
736:
730:
726:
714:
694:
675:
672:
626:
602:
598:
594:heterozygous
575:
570:
566:
564:heterozygous
559:
552:
548:
539:
535:
531:
527:
523:
519:
513:
505:
497:
486:
479:
468:
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383:
368:
336:
326:
322:
318:
312:
302:
298:
288:
284:
280:
276:
255:
240:
229:Leg striping
203:Color traits
180:
176:
175:Neither the
174:
154:plains zebra
132:
128:Chauvet Cave
113:
69:
67:
1898:Color breed
1753:Tricoloured
1730:(primarily
1590:Smoky cream
1567:Smoky black
1199:January 20,
752:melanocytes
744:outer layer
690:wild horses
421:Fjord horse
399:smoky cream
285:classic dun
249:and on the
237:Facial mask
126:such as in
1972:Categories
1732:UK English
1613:(see also
1560:Isabelline
1496:Seal brown
1268:2019-04-28
1177:2019-04-28
1105:2019-04-27
1101:. UC Davis
984:2019-04-26
900:2019-04-27
873:References
768:KIT ligand
704:expression
700:regulating
590:homozygous
586:cream gene
557:Homozygous
475:cream gene
454:Dun mimics
425:brunnblakk
375:cream gene
207:See also:
198:coloration
78:coat color
1877:Tiger eye
1781:Appaloosa
1779:See also
1605:Champagne
1494:included
1244:1651-6206
708:hair bulb
686:wild type
676:Prior to
489:blue roan
445:gulblakk.
327:mouse dun
289:zebra dun
196:wild type
116:wild type
61:A red dun
1923:KIT gene
1908:Wildtype
1872:Mushroom
1807:Rabicano
1748:Skewbald
1580:Cremello
1555:Palomino
1550:Buckskin
1479:Chestnut
1396:June 26,
1369:23844108
1329:PLOS ONE
1310:34432873
1049:26691985
960:22065780
867:non-dun1
863:non-dun2
859:non-dun2
851:Non-dun2
843:non-dun1
819:non-dun1
803:Non-dun1
795:non-dun2
791:non-dun1
766:encodes
715:non-dun2
571:non-dun2
544:dominant
536:Non-dun2
528:Non-dun1
510:Genetics
501:palomino
471:buckskin
441:palomino
437:ulsblakk
429:rødblakk
414:non-dun2
410:non-dun1
391:cremello
323:blue dun
307:chestnut
303:claybank
266:Chestnut
243:dilution
181:non-dun2
179:nor the
177:non-dun1
99:chestnut
70:dun gene
1913:Melanin
1847:Pangaré
1837:Cropout
1832:Brindle
1743:Piebald
1712:Tobiano
1585:Perlino
1360:3699485
1337:Bibcode
1126:J Hered
1040:4731265
951:3219153
928:Bibcode
839:thymine
831:guanine
815:adenine
813:has an
807:guanine
621:protein
516:alleles
482:perlino
403:alleles
395:perlino
360:donkeys
299:Red dun
281:bay dun
192:Lascaux
120:equines
1918:Agouti
1867:Flaxen
1717:Tovero
1705:Sabino
1617:Grullo
1484:Sorrel
1367:
1357:
1308:
1242:
1232:
1146:731005
1144:
1047:
1037:
958:
948:
809:where
805:has a
629:zebras
397:, and
319:grulla
314:Grullo
170:Zebras
166:tarpan
164:, the
158:quagga
156:, the
142:onager
46:grullo
1862:Sooty
1842:Liver
1820:Other
1700:Frame
1690:Overo
1639:White
1625:Pearl
1537:Cream
1474:Black
1226:(PDF)
789:Both
784:KITLG
776:KITLG
772:KITLG
764:KITLG
702:TBX3
605:T-box
567:d1/d2
553:d1/d2
549:d1/d1
449:kvit.
331:black
287:, or
262:black
162:horse
146:kiang
133:Equus
103:black
82:horse
80:of a
72:is a
1938:PAX3
1933:MITF
1802:Roan
1783:and
1520:Gray
1515:Gray
1398:2016
1365:PMID
1306:PMID
1240:ISSN
1230:ISBN
1201:2010
1142:PMID
1045:PMID
956:PMID
920:PNAS
855:TBX3
799:TBX3
793:and
780:TBX3
760:MITF
758:and
748:TBX3
738:TBX3
732:TBX5
727:TBX3
723:MSX2
721:and
719:ALX4
609:TBX3
493:gray
419:The
412:and
270:gray
211:and
86:mane
68:The
1610:Dun
1491:Bay
1355:PMC
1345:doi
1296:doi
1292:112
1134:doi
1035:PMC
1027:doi
946:PMC
936:doi
924:108
841:in
835:dun
833:in
827:dun
811:dun
756:KIT
607:3 (
551:or
520:dun
433:grĂĄ
325:or
317:or
293:bay
277:Dun
258:bay
1974::
1389:.
1377:^
1363:.
1353:.
1343:.
1331:.
1327:.
1304:.
1290:.
1286:.
1260:.
1238:.
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1140:.
1130:69
1128:.
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1088:^
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1021:.
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993:^
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954:.
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881:^
869:.
801:.
754:,
631:.
611:)
540:d2
532:d1
393:,
283:,
168:.
148:,
144:,
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1450:e
1443:t
1436:v
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1136::
1108:.
1051:.
1029::
987:.
962:.
938::
930::
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538:(
530:(
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522:(
389:(
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.