OMIA 000209-9796 : Coat colour, dominant white in Equus caballus

See the equivalent entry at NCBI

In other species: rabbit , pig , cattle , dog , domestic cat , llama , alpaca , Arctic fox , ass , raccoon dog

Possible human homologue (MIM number): 172800

Mendelian trait/disorder: yes

Mode of inheritance: Autosomal Dominant

Considered a defect: no

Key mutation known: yes

Year key mutation first reported: 2005

Cross-species summary: The dominant white gene is one of a number of genes that regulate normal growth and proliferation of cells. In fact, it encodes a protein that protrudes through the cell membrane, relaying 'messages' across the membrane, from outside to inside the cell. The transmembrane domain of the protein is a receptor for a growth factor (a protein produced by one type of cell, that acts on another type of cell). The domain inside the cell has tyrosine kinase activity. When a growth factor binds to the receptor on the outside of the cell, this stimulates tyrosine kinase activity inside the cell, which sets off a cascade of phosphorylations, resulting in activation of transcription factors, which in turn activate genes, resulting in multiplication of stem cells, including melanocyte precursor cells, in the developing embryo. This whole process is known as a signal transduction pathway. During embryonic development, the melanosome precursor cells migrate from the neural crest down either side of the body. Under normal circumstances, they eventually meet at the centre of the belly. The cells then proliferate in all directions until they meeting neighbouring cells, thereby filling up all available areas, resulting in a solid mass of melanocytes over the entire body. The dominant white allele produces a defective transmembrane protein which is unable to relay 'messages', resulting in a lack of melanocytes, and hence white coat colour. An interesting aspect of the dominant white gene is that if it is activated at the wrong time, the result can be excess and uncontrolled proliferation of stem cells; in other words, cancer. In fact, at some time in the past, a feline retrovirus (the Hardy-Zuckerman 4 feline sarcoma virus) 'picked up' (by transduction) a copy of the dominant white gene from a cat, and incorporated this gene into its own genome. When this retrovirus infects cats, it activates its own copy of the gene at inappropriate times, causing sarcoma - a malignant tumour of cells derived from connective tissue. Retroviral genes that cause cancer are called oncogenes. The original host version of an oncogene is called a proto-oncogene. Thus, the dominant white gene is actually a proto-oncogene. In this particular case, the oncogene was discovered and named v-kit (where 'v' indicates a viral version of the gene) long before its association with white coat colour was established. The corresponding proto-oncogene is called c-kit, where 'c' stands for cellular. After the discovery and cloning of v-kit in the feline retrovirus by Besmer et al. (1986; Nature 320:415-421), c-kit was identified and mapped first in humans, by Mattei et al. (1987; Cytogenetics and Cell Genetics 46:657 only), and then in mice (Chabot et al., 1988; Nature 335:88-89, 1988), where it was shown to be identical with the long-recognised white-spotting (W) locus. Three years later, Giebel and Spritz (1991; Proceedings of the National Academy of Sciences 88:8696-8699) showed that mutations at the c-kit gene in humans cause piebaldism, which is the human homologue of white spotting (see the MIM entry at the top of this page)

Species-specific name: Tobiano, Sabino

Mapping: Mau et al. (2004) mapped this trait to a region of chromosome ECA3 containing the KIT gene.

Molecular basis: Using the comparative candidate-gene strategy (based on the KIT gene being associated with similar coat-colour-phenotypes in humans and pigs), Brooks and Bailey (2005) sequenced the KIT gene in horses of each of the three genotypes at a Sabino-spotting locus they called Sabino 1, and identified a splice variant, namely "a base substitution for T with A in intron 16, 1037 bases following exon 16" that results in the skipping of exon 17 and which was entirely associated with the SB1 allele. Homozygosity for this allele "results in a complete or nearly completely white phenotype". Haase et al. (2007) showed that the dominant white coat colour is due to a range of mutations in the KIT gene across a range of breeds. Brooks et al. (2007) showed that the tobiano pattern is due to a large paracentric inversion that begins just downstream of the KIT gene. In the words of Brooks et al. (2007), the "inversion may disrupt regulatory sequences for the KIT gene and cause the white spotting pattern". Haase et al. (2008) showed that the same inversion is the cause of this coat colour in a range of German horse breeds, indicating that it is an old mutation. Additional KIT mutations associated with white or white spotting have been reported by Haase et al. (2009), Holl et al. (2010), Haase et al. (2011) and Hauswirth et al. (2013). To help keep track of the large number of apparently functional mutations, Haase et al. (2009) proposed a KIT-allele naming system (W1, . . . , W11) for the 11 mutants then recognised. The new allele reported by Holl et al. (2010) is W12. Haase et al. (2011) reported new alleles W13-W17. The three new alleles reported by Hauswirth et al. (2013) are named W18, W19, and W20. Another new allele, W21, was reported by Haase et al. (2015). W22 was reported by Dürig et al. (2017) and W23 by Holl et al. (2017), who also provided the first report of a viable W15 homozygote, which was completely white, the first such reported case of viable homozygosity for any W allele.

Table S1 of Dürig et al. (2017) provides details of all KIT alleles (plus alleles at PAX3, MITF and EDNRB) known in April 2017.

Negro et al. (2017): "a missense mutation (p.Arg682His) in KIT [= allele W20] was associated with white facial markings (P < 0.05) and with total white markings (P < 0.05) in PRMe horses. The relative contribution of this variant to white markings in PRMe horses was estimated at 47.6% (head) and 43.4% (total score). In PRE horses, this variant was also associated with hindlimb scores (P < 0.05) with a relative contribution of 41.2%."

Capomaccio et al. (2017) reported a de novo likely causal (splicing) variant (NC_009146.2:g.77736559C>T, called allele W24) in a white Italian Trotter born to two non-white (bay) parents.

Hoban et al. (2018) reported three new likely causal variants, which they designated "W25 (p.Leu223Pro) [in a Thoroughbred family], W26 (p.Ser846Valfs*15) [in the same Thoroughbred family] and W27 (p.Cys491Trp) [in a second Thoroughbred family]". These authors also identified variant W13 in a Miniature Horse family.

Prevalence: Druml et al. (2018) reported that the SB1 variant is "present in three breeds (Haflinger, 14 out of 98; Noriker, four out of 189; Lipizzan one out of 329) . . . None of the SB1/sb1-carrier horses met the criteria defining the Sabino1 pattern according to current applied protocols."

Associated gene:

Symbol Description Species Chr Location OMIA gene details page Other Links
KIT v-kit Hardy-Zuckerman 4 feline sarcoma viral oncogene homolog Equus caballus 3 NC_009146.3 (79618685..79536564) KIT Homologene, Ensembl, NCBI gene

Variants

By default, variants are sorted chronologically by year of publication, to provide a historical perspective.

Readers can re-sort on any column by clicking on the column header. Click it again to sort in a descending order. To create a multiple-field sort, hold down Shift while clicking on the required column headers

Breed(s) Variant Phenotype Gene Allele Type of Variant Reference Sequence Chr. g. or m. c. or n. p. Verbal Description EVA ID Year Published PubMed ID(s) Acknowledgements
Thoroughbred Coat colour, dominant white KIT W27 missense 3 c.1473T>G p.Cys491Trp 2018 29333746
Thoroughbred Coat colour, dominant white KIT W26 deletion, small (<=20) 3 c.2536delA p.Ser846Valfs*15 2018 29333746
South German Draft Coat colour, dominant white KIT W11 splicing Equ cab 2 3 g.77731743G>A c.2684+1G>A 2009 19456317 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Miniature Horse Quarter Horse Coat colour, dominant white KIT W13 splicing Equ cab 2 3 g.77735380G>C c.2472+5G>C 2011 21554354 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Appaloosa Haflinger Lipizzan Noriker Quarter Horse Coat colour, dominant white KIT sabino 1 (SB1) splicing Equ cab 2 3 g.77735520A>T "SNP AX-103727726 (ECA3:77735520; SB1, SNP KI16+1037A)" (Druml et al., 2018) 2005 16284805 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Icelandic Coat colour, dominant white KIT W8 splicing Equ cab 2 3 g.77736688 c.2222-1 2009 19456317 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Swiss Warmblood Coat colour, dominant white KIT W18 splicing Equ cab 2 3 g.77745065G>A c.1346 + 1G>A 2013 23659293 Variant coordinates obtained from or confirmed by EBI's Some Effect Predictor (VEP) tool; breed and allele information obtained from Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Thoroughbred Coat colour, dominant white KIT W7 splicing Equ cab 2 3 g.77771305G>C c.338-1G>C 2009 19456317 Variant coordinates obtained from or confirmed by EBI's Some Effect Predictor (VEP) tool; breed and allele information obtained from Table S1 of Durig et al. (2017) Anim Genet 48:123-124
White spotting KIT W22 deletion, gross (>20) EquCab2 3 g.77,740,239_77,742,136del1898 2017 28444912
Oldenburg Coat colour, dominant white KIT W16 missense EquCab2 3 g.77732055A>T c.2489A>T p.K830I 2011 21554354 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Thoroughbred Coat colour, dominant white KIT W14 deletion, gross (>20) EquCab2 3 g.77735465_77735518del c.2392_2445del p.H798_N815del 2011 21554354 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Icelandic White spotting KIT W21 deletion, small (<=20) EquCab2 3 g.77735488delG c.2369delC p.Ala790Glufs*20 2015 26059442 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Thoroughbred Coat colour, dominant white KIT W5 deletion, small (<=20) EquCab2 3 g.77737214delG c.2193delG p.T732QfsX9 2009 19456317 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Franches-Montagnes Coat colour, dominant white KIT W1 nonsense (stop-gain) EquCab2 3 g.77737256C>G c.2151C>G p.Y717* 2007 17997609 Variant coordinates obtained from or confirmed by EBI's Some Effect Predictor (VEP) tool; breed and allele information obtained from Table S1 of Durig et al. (2017) Anim Genet 48:123-124
American Paint Horse Appaloosa German Riding Pony Gipsy Noriker Old-Tori Oldenberg Quarter Horse Thoroughbred Warmblood Welsh Pony Coat colour, dominant white KIT W20 missense EquCab2 3 g.77739534G>A c.2045G>A p.R682H 2013 23659293 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Japanese Draft Coat colour, dominant white KIT W17b missense EquCab2 3 g.77739558T>C c.2021T>C p.L674P 2011 21554354 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Japanese Draft Coat colour, dominant white KIT w17a missense EquCab2 3 g.77739579A>T c.2001A>T p.E667D 2011 21554354 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Thoroughbred Coat colour, dominant white KIT W2 missense EquCab2 3 g.77740854G>A c.1960G>A p.G654R 2007 17997609 Variant coordinates obtained from or confirmed by EBI's Some Effect Predictor (VEP) tool; breed and allele information obtained from Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Camarillo White Horse Coat colour, dominant white KIT W4 missense EquCab2 3 g.77741094C>T c.1805C>T p.A602V 2007 17997609 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Holstein Coat colour, dominant white KIT W9 missense EquCab2 3 g.77741111G>A c.1789G>A p.G597R 2009 19456317 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Arabian Coat colour, dominant white KIT W15 missense EquCab2 3 g.77741665T>C c.1597T>C p.C533R 2011 21554354 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Arabian Coat colour, dominant white KIT W19 missense EquCab2 3 g.77745090A>G c.1322A>G p.Y441C 2013 23659293 Variant coordinates obtained from or confirmed by EBI's Some Effect Predictor (VEP) tool; breed and allele information obtained from Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Quarter Horse Coat colour, dominant white KIT W10 deletion, small (<=20) EquCab2 3 g.77758240_77758243del c.1126_1129delGAAC p.E376FfsX3 2009 19456317 Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Thoroughbred Coat colour, dominant white KIT W6 missense EquCab2 3 g.77765059G>A c.856G>A p.G286R 2009 19456317 Variant coordinates obtained from or confirmed by EBI's Some Effect Predictor (VEP) tool; breed and allele information obtained from Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Arabian White spotting KIT W23 splicing EquCab2 3 g.77769789C>G c.756+1G>C ss2137493881 2017 28378922
Arabian Coat colour, dominant white KIT W3 nonsense (stop-gain) EquCab2 3 g.77769840A>T c.706A>T p.K236* 2007 17997609 Variant coordinates obtained from or confirmed by EBI's Some Effect Predictor (VEP) tool; breed and allele information obtained from Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Thoroughbred Coat colour, dominant white KIT W12 deletion, small (<=20) EquCab2 3 g.77771079_77771083delTCTGC c.559_563delTCTGC p.Ser187ArgfsX10 2010 Reference not in PubMed; see OMIA 000209-9796 for reference details Variant coordinates gleaned from or confirmed by Table S1 of Durig et al. (2017) Anim Genet 48:123-124
Italian Trotter Coat colour, dominant white KIT W24 splicing EquCab2.0 3 g.77736559C>T 2017 28856698
Thoroughbred Coat colour, dominant white KIT W25 missense EquCab2.0 3 g.77769878A>G c.668T>C p.Leu223Pro 2018 29333746

References


Note: the references are listed in reverse chronological order (from the most recent year to the earliest year), and alphabetically by first author within a year.
2018 Druml, T., Grilz-Seger, G., Neuditschko, M., Horna, M., Ricard, A., Pausch, H., Brem, G. :
Novel insights into Sabino1 and splashed white coat color patterns in horses. Anim Genet 49:249-253, 2018. Pubmed reference: 29635692. DOI: 10.1111/age.12657.
Hoban, R., Castle, K., Hamilton, N., Haase, B. :
Novel KIT variants for dominant white in the Australian horse population. Anim Genet :, 2018. Pubmed reference: 29333746. DOI: 10.1111/age.12627.
Li, B., He, X., Zhao, Y., Bai, D., Shiraigo, W., Zhao, Q., Manglai, D. :
Regulatory pathway analysis of coat color genes in Mongolian horses. Hereditas 155:13, 2018. Pubmed reference: 28974924. DOI: 10.1186/s41065-017-0048-y.
2017 Capomaccio, S., Milanesi, M., Nocelli, C., Giontella, A., Verini-Supplizi, A., Branca, M., Silvestrelli, M., Cappelli, K. :
Splicing site disruption in the KIT gene as strong candidate for white dominant phenotype in an Italian Trotter. Anim Genet 48:727-728, 2017. Pubmed reference: 28856698. DOI: 10.1111/age.12590.
Dürig, N., Jude, R., Holl, H., Brooks, S.A., Lafayette, C., Jagannathan, V., Leeb, T. :
Whole genome sequencing reveals a novel deletion variant in the KIT gene in horses with white spotted coat colour phenotypes. Anim Genet 48:483-485, 2017. Pubmed reference: 28444912. DOI: 10.1111/age.12556.
Holl, H.M., Brooks, S.A., Carpenter, M.L., Bustamante, C.D., Lafayette, C. :
A novel splice mutation within equine KIT and the W15 allele in the homozygous state lead to all white coat color phenotypes. Anim Genet 48:497-498, 2017. Pubmed reference: 28378922. DOI: 10.1111/age.12554.
Kim, N.Y., Bhuiyan, M.S.A., Chae, H.S., Baek, K.S., Son, J.K., Shin, S.M., Woo, J.H., Park, S.H., Lee, S.H. :
Genome-wide association study for tobiano spotting coat color in Korean Jeju × Thoroughbred horse population. Anim Genet 48:728-729, 2017. Pubmed reference: 28850680. DOI: 10.1111/age.12596.
Negro, S., Imsland, F., Valera, M., Molina, A., Solé, M., Andersson, L. :
Association analysis of KIT, MITF, and PAX3 variants with white markings in Spanish horses. Anim Genet 48:349-352, 2017. Pubmed reference: 28084638. DOI: 10.1111/age.12528.
2016 Han, H., Mao, C., Chen, N., Lan, X., Chen, H., Lei, C., Dang, R. :
Single nucleotide polymorphisms of Kit gene in Chinese indigenous horses. Jpn J Vet Res 64:81-9, 2016. Pubmed reference: 27348891.
Martin, L., Damaso, N., Mills, D. :
Detection of single nucleotide polymorphisms (SNP) in equine coat color genes using SNaPshot(TM) Multiplex kit or Pluronic F-108 tri-block copolymer and capillary electrophoresis. Electrophoresis :, 2016. Pubmed reference: 27542347. DOI: 10.1002/elps.201600245.
Reissmann, M., Musa, L., Zakizadeh, S., Ludwig, A. :
Distribution of coat-color-associated alleles in the domestic horse population and Przewalski's horse. J Appl Genet 57:519-525, 2016. Pubmed reference: 27194311. DOI: 10.1007/s13353-016-0352-7.
2015 Haase, B., Jagannathan, V., Rieder, S., Leeb, T. :
A novel KIT variant in an Icelandic horse with white-spotted coat colour. Anim Genet 46:466, 2015. Pubmed reference: 26059442. DOI: 10.1111/age.12313.
2013 Hauswirth, R., Jude, R., Haase, B., Bellone, R.R., Archer, S., Holl, H., Brooks, S.A., Tozaki, T., Penedo, M.C., Rieder, S., Leeb, T. :
Novel variants in the KIT and PAX3 genes in horses with white-spotted coat colour phenotypes. Anim Genet 44:763-5, 2013. Pubmed reference: 23659293. DOI: 10.1111/age.12057.
2011 Bai, D.Y., Yang, L.H., Unerhu, U., Zhao, Y.P., Zhao, Q.N., Hasigaowa, H., Dugarjaviin, M. :
[Effects of Kit gene on coat depigmentation in white horses]. Yi Chuan 33:1171-8, 2011. Pubmed reference: 22120071.
Haase, B., Rieder, S., Tozaki, T., Hasegawa, T., Penedo, MC., Jude, R., Leeb, T. :
Five novel KIT mutations in horses with white coat colour phenotypes. Anim Genet 42:337-339, 2011. Pubmed reference: 21554354. DOI: 10.1111/j.1365-2052.2011.02173.x.
2010 Bellone, RR. :
Pleiotropic effects of pigmentation genes in horses. Anim Genet 41 Suppl 2:100-10, 2010. Pubmed reference: 21070283. DOI: 10.1111/j.1365-2052.2010.02116.x.
Holl, H., Brooks, S., Bailey, E. :
De novo mutation of KIT discovered as a result of a non-hereditary white coat colour pattern Animal Genetics 41 Suppl 2:196-198, 2010.
Horváth, G., Blahó, M., Kriska, G., Hegedüs, R., Gerics, B., Farkas, R., Akesson, S. :
An unexpected advantage of whiteness in horses: the most horsefly-proof horse has a depolarizing white coat. Proc Biol Sci 277:1643-50, 2010. Pubmed reference: 20129982. DOI: 10.1098/rspb.2009.2202.
Li, B., He, XL., Zhao, YP., Wang, XJ., Manglai, D., Zhang, YR. :
[Molecular basis and applicability in equine color genetics]. Yi Chuan 32:1133-40, 2010. Pubmed reference: 21513164.
2009 Haase, B., Brooks, SA., Tozaki, T., Burger, D., Poncet, PA., Rieder, S., Hasegawa, T., Penedo, C., Leeb, T. :
Seven novel KIT mutations in horses with white coat colour phenotypes. Anim Genet 40:623-629, 2009. Pubmed reference: 19456317. DOI: 10.1111/j.1365-2052.2009.01893.x.
Haase, B., Obexer-Ruff, G., Dolf, G., Rieder, S., Burger, D., Poncet, PA., Gerber, V., Howard, J., Leeb, T. :
Haematological parameters are normal in dominant white Franches-Montagnes horses carrying a KIT mutation. Vet J :, 2009. Pubmed reference: 19362501. DOI: 10.1016/j.tvjl.2009.02.017.
Kakoi, H., Tozaki, T., Nagata, S., Gawahara, H., Kijima-Suda, I. :
Development of a method for simultaneously genotyping multiple horse coat colour loci and genetic investigation of basic colour variation in Thoroughbred and Misaki horses in Japan. J Anim Breed Genet 126:425-31, 2009. Pubmed reference: 19912416. DOI: 10.1111/j.1439-0388.2009.00841.x.
Lewin, H.A. :
Genetics. It's a bull's market. Science 324:478-9, 2009. Pubmed reference: 19390037. DOI: 10.1126/science.1173880.
Rieder, S. :
Molecular tests for coat colours in horses. J Anim Breed Genet 126:415-24, 2009. Pubmed reference: 19912415. DOI: 10.1111/j.1439-0388.2009.00832.x.
2008 Haase, B., Jude, R., Brooks, SA., Leeb, T. :
An equine chromosome 3 inversion is associated with the tobiano spotting pattern in German horse breeds. Anim Genet 39:306-9, 2008. Pubmed reference: 18410476. DOI: 10.1111/j.1365-2052.2008.01715.x.
Rieder, S., Hagger, C., Obexer-Ruff, G., Leeb, T., Poncet, PA. :
Genetic analysis of white facial and leg markings in the Swiss Franches-Montagnes Horse Breed. J Hered 99:130-6, 2008. Pubmed reference: 18296388. DOI: 10.1093/jhered/esm115.
2007 Brooks, SA., Lear, TL., Adelson, DL., Bailey, E. :
A chromosome inversion near the KIT gene and the Tobiano spotting pattern in horses. Cytogenet Genome Res 119:225-30, 2007. Pubmed reference: 18253033. DOI: 10.1159/000112065.
Haase, B., Brooks, SA., Schlumbaum, A., Azor, PJ., Bailey, E., Alaeddine, F., Mevissen, M., Burger, D., Poncet, PA., Rieder, S., Leeb, T. :
Allelic heterogeneity at the equine KIT locus in dominant white (W) horses. PLoS Genet 3:e195, 2007. Pubmed reference: 17997609. DOI: 10.1371/journal.pgen.0030195.
2005 Brooks, SA., Bailey, E. :
Exon skipping in the KIT gene causes a Sabino spotting pattern in horses. Mamm Genome 16:893-902, 2005. Pubmed reference: 16284805. DOI: 10.1007/s00335-005-2472-y.
2004 Mau, C,, Poncet, P.A., Bucher, B., Stranzinger, G., Rieder, S, :
Genetic mapping of dominant white (W), a homozygous lethal condition in the horse (Equus caballus) Journal of Animal Breeding and Genetics 121:374-383, 2004.
2002 Brooks, S.A., Terry, R.B., Bailey, E. :
A PCR-RFLP for KIT associated with tobiano spotting pattern in horses Animal Genetics 33:301-303, 2002. Pubmed reference: 12139510.
1978 Trommershausen-Smith, A. :
Linkage of tobiano coat spotting and albumin markers in a pony family Journal of Heredity 69:214-216, 1978. Pubmed reference: 569673.
1969 Pulos, WL., Hutt, FB. :
Lethal dominant white in horses. J Hered 60:59-63, 1969. Pubmed reference: 5816567.

Edit History


  • Created by Kao Castle on 03 Aug 2009
  • Changed by Frank Nicholas on 11 Sep 2011
  • Changed by Frank Nicholas on 14 Sep 2011
  • Changed by Frank Nicholas on 09 Dec 2011
  • Changed by Frank Nicholas on 06 Jan 2013
  • Changed by Frank Nicholas on 13 May 2013
  • Changed by Frank Nicholas on 29 Nov 2013
  • Changed by Tosso Leeb on 07 Aug 2015
  • Changed by Frank Nicholas on 24 Aug 2017
  • Changed by Frank Nicholas on 01 Sep 2017
  • Changed by Frank Nicholas on 24 Nov 2017
  • Changed by Frank Nicholas on 19 Jan 2018
  • Changed by Frank Nicholas on 19 Apr 2018