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Dun gene

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649: 237: 470: 661: 677: 67: 245: 229: 1427: 1518: 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. 50: 198: 36: 634:
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
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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
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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
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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.
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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
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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.
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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. 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.
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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."
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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.
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Cieslak, Jakub; Brooks, Samantha Ann; Wodas, Lukasz; Mantaj, Weronika; Borowska, Alicja; Sliwowska, Joanna Helena; Ziarniak, Kamil; MacKowski, Mariusz (2021).
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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.
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breed, which is predominantly dun, uses unique Norwegian-based terminology to distinguish between the different shades of dun horses. "Brown dun", or
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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
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Historically, before modern genetic studies distinguished between alleles, diluted colors were sometimes lumped together and simply called "dun".
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deletion is a more derived allele. Nucleotide diversity across the flanking regions of chromosome 8 for the various alleles indicates that the
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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
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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: 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 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 1938: 1244: 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
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that also carries dun, showing primitive dorsal striping or leg bars indicative of a red dun may be called a "dunalino."
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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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at chromosome 8 base pair 18,226,905, which appears to be sufficient to cause non-dun1 coloration. In addition,
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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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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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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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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
604: 574: 545:) horses do not have dun dilution but may exhibit some primitive markings. 499: 164: 138: 96: 1068:"A horse of a different color: Genetics of camouflage and the dun pattern" 1908: 1872: 1688: 1577: 1501: 1484: 1233:
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.).
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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
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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
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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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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
619: 446:- literally "gray" - is a grullo, buckskin duns are called 197: 95:. The dun gene lightens most of the body while leaving the 49: 1023: 907:. Veterinary Genetics Lab, University of California, Davis 365:
Transverse stripe, a crosswise stripe along the shoulders
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are dun, and several closely related species in the genus
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A cremello, perlino or smoky cream is called "white" or
35: 1333: 1398:"Genetics of camouflage and the Dun pattern in horses" 793:
was expressed. TBX3 is not thought to directly affect
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horses typically have clearer primitive markings than
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in that it acts on any coat color. In contrast, the
1130: 1128: 1126: 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 1980: 1225: 1223: 1221: 1134: 1123: 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, 1104: 1102: 1100: 1087: 1085: 1083: 1453: 1218: 1019: 1017: 1015: 1013: 1011: 1009: 1007: 1005: 537:), produces dilution and primitive markings. 899: 897: 895: 893: 412:). Thus, if a horse with two cream dilution 388:(single copy) = "dunalino" or "palomino dun" 147:show dun characteristics. These include the 1097: 1080: 724:during hair growth." The region deleted in 348:Another characteristic of the dun gene are 1460: 1446: 1391: 1389: 1195: 1002: 918: 194:mutations were found in any other equids. 1369: 1359: 1310: 1049: 960: 950: 890: 120: 675: 599:is an incomplete dominant which must be 595:acts only on black-based coats, and the 468: 243: 235: 227: 196: 1386: 1229: 1182:"More about Dun and Primitive Markings" 924: 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 14: 1981: 1467: 1201: 491:A red dun may also be confused with a 450:or white dun, and a "dunalino" (dun + 1441: 1395: 1174: 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 1546: 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. 24: 671: 25: 2005: 1419: 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: 1327: 1286: 1261: 1239:(Thesis). Uppsala. p. 42. 860:, so is not necessary for dun. 397:cream gene on a black base coat 213: 171:, and an extinct subspecies of 1402:SciLifeLab, Uppsala University 1163: 977: 834:single nucleotide polymorphism 454:) is called a "yellow dun" or 271:(bay, classic, or zebra dun), 13: 1: 883: 695:are thought to have been the 464: 1361:10.1371/journal.pone.0067841 905:"Dun Dilution - Direct Test" 7: 836:compared to the version of 584:lacks primitive markings. 520: 275:(mouse dun or grullo), and 163:, an extinct subspecies of 10: 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: 1896: 1830: 1805: 1776: 1736: 1696: 1687: 1674: 1648: 1586: 1555: 1537: 1525: 1514: 1475: 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: 234: 224:Primitive markings 211: 149:Przewalski's horse 101:primitive markings 99:, tail, legs, and 1989:Horse coat colors 1976: 1975: 1826: 1825: 1772: 1771: 1612: 1611: 1608: 1607: 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 1746: 1694: 1693: 1685: 1684: 1632: 1574: 1553: 1552: 1544: 1543: 1520: 1509: 1498: 1477:Base coat colors 1462: 1455: 1448: 1439: 1438: 1429: 1413: 1412: 1410: 1408: 1393: 1384: 1383: 1373: 1363: 1331: 1325: 1324: 1314: 1290: 1284: 1283: 1281: 1280: 1275:. 15 August 2017 1273:Cowgirl Magazine 1265: 1259: 1258: 1238: 1227: 1216: 1215: 1213: 1211: 1199: 1193: 1192: 1190: 1189: 1178: 1172: 1167: 1161: 1160: 1132: 1121: 1120: 1118: 1117: 1106: 1095: 1089: 1078: 1075: 1063: 1053: 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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: 476: 459: 455: 447: 443: 439: 435: 429: 424: 420: 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: 18:Dun (colour) 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:PangarĂ© 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 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:. 1044:. 1034:48 1032:. 1028:. 1004:^ 987:. 965:. 955:. 945:. 933:. 929:. 892:^ 880:. 812:. 765:, 642:. 622:) 551:d2 543:d1 404:, 294:, 179:. 159:, 155:, 151:, 1745:) 1631:) 1573:) 1497:) 1461:e 1454:t 1447:v 1411:. 1382:. 1358:: 1350:: 1344:8 1323:. 1309:: 1282:. 1257:. 1214:. 1191:. 1159:. 1147:: 1119:. 1062:. 1040:: 998:. 973:. 949:: 941:: 914:. 549:( 541:( 535:D 533:( 400:( 20:)

Index

Dun (colour)


grullo

dilution gene
coat color
horse
mane
primitive markings
coat color genetics
chestnut
black
wild type
equines
cave paintings
Chauvet Cave
Equus
Przewalski's horse
onager
kiang
African wild ass
plains zebra
quagga
horse
tarpan
Zebras

Lascaux
wild type

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