- Email: [email protected]
On the probable genotype of domestic cats in ancient Egypt
Journal
of Archaeological
Science 1982, 9, 377-379
On the Probable Genotype of Domestic Cats in Ancient Egypt Bennett Blumenberg” Genetic and archaeological evidence suggests that the majority of mummified ritual cats (Felis Kbyca subsp.) in ancient Egypt carried the t + allele and exhibited the striped tabby phenotype. A few of these ritual cats, however, appear to be F. chum subsp., most likely carried the T” allele and were characterized by the Abyssinian phenotype. Severaltypes of evidence suggest that the T”allele first became fixed in a Felis sp. population in either southeast Asia or northeast Africa. A number of Felis subspecies appear to be plausible candidates for the population of T” origin. Keywords: ABYSSINIAN, ALLELE, ANCIENT EGYPT, CHINA, FELZS CATUS, FELZS CHAUS, FELZS LZBYCA OCREATA, GENOTYPE, ALLELE,
PHENOTYPE, POPULATION
GENETICS, STRIPED TABBY, THAILAND.
The paper by Armitage & Clutton-Brock (1981) is a most valuable contribution with respect to (I) the anatomical and histological data established; (2) the elucidation of mummification techniques; and (3) the probable role that domestic cats played in the ritual life of ancient Egyptians. Additional observations that may be of interest can be offered concerning the genotype of these cats. On p. 192, the authors report that microscopic examination established the agouti character of the cat hair. They conclude from this observation that the overall phenotype of these Egyptian cats would have been either striped tabby or Abyssinian. Recent genetic research has established that these two phenotypes are mutually exclusive in Felis sp., for they are controlled by different alleles of the tabby gene (Robinson, 1977). There are three well known forms (alleles) of this gene and they produce the phenotypes known as Abyssinian, striped tabby and blotched tabby. The dominance relationships between these alleles are T” > t f > tb (Robinson, 1977). The t + allele produces the phenotype known as striped tabby that is characterized by vertical curving stripes, short bars or spots on the sides of the body. The head, legs and tail are also distinctively striped; see figure 1 in Robinson (1977). This pattern is believed to represent the ancestral or “wild type” phenotype that is diagnostic of the population (s?) representing the reservoir(s) from which the first domestic groups were drawn (Todd, 1978). Early domestication certainly occurred in Egypt (Armitage & Clutton-Brock, 1981; Baldwin, 1975) and quite possibly in the Indus Valley as well (Ahmad et al., 1980). The Ta allele produces the Abyssinian phenotype in which the striped pattern may be restricted to the head, although frequently found on the legs and tail; see figure 3 in Robinson (1977). While the name “Abyssinian” suggests an origin in northeast Africa,
0305~4403/82/040377+03
“Faculty of Sciences, Lesley College, 29 Everett St, Cambridge, Mass. 02238, U.S.A. 377 $03.00/O 0 1982 Academic Press Inc. (London) Limited
378
B. BLUMENBERG
there is no evidence that this mutant was common in ancient Egypt. Egyptian cat populations now exhibit a genetic profile that is best described as provincial European and the Abyssinian allele is extremely rare or nonexistent (Todd & Blumenberg, 1978). The highest known frequencies of the T” allele ( > 10 %) are found in the isolated Thai city of Mai Hong Son, Hong Kong, Macau and Singapore (Fagen & Meeswat, 1981; Searle, 1959). The likelihood that the Abyssinian allele was present at very low frequencies in the gene pool of early domestic cats in Egypt supports the observation that F. chaus represents very few of those ritual cats that were eventually mummified (Armitage & Clutton-Brock, 1981; Morrison-Scott, 1952). These few F. chaus are significant, however, when hypotheses concerning the origin of the Ta allele are considered (see below). This species is characterized by a phenotype that is essentially devoid of body pattern (Pocock, 1951). F. libyca libyca, which frequently exhibits a well defined tabby pattern (Pocock, 1951, pp. 61-65), appears to represent those populations that quickly came to dominate the domestication process, be this phenotype given distinct species status (Pocock, 1951), believed conspecific with F. silvestris (Todd, 1978) or assigned subspecies rank as F. silvestris libyca (Armitage & Clutton-Brock, 1981); see figures 2 and 3 in Armitage & Clutton-Brock (198 1). Two hypotheses may be offered concerning plausible geographic localities in which the Ta allele first arose and became fixed in Felis sp. populations. Hypothesis 1 suggests a southeast or east Asian origin for this mutant allele and is supported by the biogeographic genetic evidence cited above. F. chaus jiilvidina is found in Burma, Thailand and Vietnam and is characterized by a phenotype virtually devoid of body pattern (Pocock, 1951, pp. 175-176). Pocock (1951, pp. 13-14) describes several feral domestic cat (F. catus) specimens from Burma whose body patterns were faint and obscure. Gray (1972, p. 36) reports that interspecific matings between F. chaus (subspecies not identified) and F. cam produced fertile F, hybrids; such could be the mechanism by which the Ta allele entered the F. catus gene pool. A less attractive variation of this hypothesis could propose that F. bieti bieti, whose “body stripes are sometimes obliterated” (Pocock, 195 1, p. 134) and whose range is the western Chinese provinces of Szechwan and Kansu (now Xinjiang and Xizang), provided the populations of origin for the T” allele. Hypothesis 2 develops from a consideration of the evidence available from Egyptian archaeozoology (Armitage & Clutton-Brock, 1981; Baldwin, 1975; Morrison-Scott, 1952) that points to a northeast African origin for the T” allele. All of these authors agree that a few of the Egyptian ritual cats that were mummified appear to be F. chaus. F. chaus nilotica of Egypt represents one possible source for the T” allele (Flower, 1932, p. 390; Gray, 1972; Pocock, 1951, p. 159). However, it is not necessary to postulate F. chaus as the species in which the T” allele first arose. F. libyca ocreata is essentially devoid of body pattern and has long been known from the Ethiopian region (Allen, 1954, p. 238; Pocock, 1944, pp. 67-68, 1951, pp. 69-72; Thomas & Hinton, 1923, p. 253). Pocock (1907a, b) suggested that the domestic cat (i.e. F. catus) Abyssinian phenotype represents a genetic heritage acquired from F. ocreata (i.e. F. libyca ocreata). Kratochvil & Kratochvil (1976, p. 205) note that F. Zibyca ocreata represents a reasonable alternative to F. libyca libyca as the ancestor of the first domestic cats. Pocock (1907c) reports upon presumed hybrids between F. catus and F. libyca ocreata. If additional evidence serves to firmly establish hypothesis 2, then the origin of the name “Abyssinian” for this particular phenotype will finally have been clarified. Acknowledgements
The author would like to thank Dr J. Clutton-Brock whose comments brought the Gray (1972) reference to his attention and served to stimulate the formulation of origin hypotheses for the T” allele.
GENOTYPE OF DOMESTIC CATS
379
References Ahmad, M., Blumenbcrg, B. & Chaudhary, M. F. (1980). Mutant allele frequencies and genetic distance in cat populations of Pakistan and Asia. Journal of Heredity 71, 323-330. Allen, G. M. (1954). A checklist of African mammals. Bulletin of the Museum of Comparafive Zoology 83, l-763. Armitage, P. L. & Clutton-Brock, J. (1981). A radiological and histological investigation into the mummification of cats from ancient Egypt. Journal of Archaeological Science 8, 185-196. Baldwin, J. A. (1975). Notes and speculations on the domestication of the cat in Egypt. Anrhropos
70, 428-448.
Fagen, R. & Meeswat, K. (1981). Population genetics and demography of the domestic cat (Felis catus L. : Carnivora Felidae) in Thailand. Natural History Bulletin of the Siam Society 29, 7137-150. Flower, S. S. (1932). Notes on the recent mammals of Egypt, with a list of the species recorded from that Kingdom. Proceedings of the Zoological Society of London, pp. 369-450. Gray, A. P. (1972). Mammalian Hybrids: a Checklist with Bibliography. Farnham Royal: Commonwealth Agricultural Bureaux. Kratochvil, J. & Kratochvil, Z. (1976). The origin of domesticated forms of the genus Felis (Mammalia). Zoologicke’ Listy 25, 193-208. Morrison-Scott, T. C. S. (1952). The mummified cats of Ancient Egypt. Proceedings of the Zoological Society of London 121, 861-867. Pocock, R. I. (1907a). On the characters and names of the two types of domestic cat and the name of the European wild cat. Proceedings of rhe Zoological Society of London, pp. 149-155. Pocock, R. I. (19076). On Manx, Persian, Siamese, Indian and Abyssinian breeds. Proceedings of the Zoological Society of London, pp. 161-165. Pocock, R. I. (1907~). On alleged casesof interbreeding between domestic cats and various wild speciesof the genus Felis. Proceedings of the Zoological Society of London, pp. 165-168. Pocock, R. I. (1944). The races of the North African Wild Cat (Felis Lybica). Proceedings of the Zoological Society of London 114, 65-73. Pocock, R. I. (1951). Catalogue of the Genus Felis. London: British Museum. Robinson, R. (1977). Genetics for Car Breeders, 2nd Edn. New York: Pergamon Press. Searle, A. G. (1959). A study of variation in Singapore cats. Journal of Genetics 56, 1-17. Thomas, 0. & Hinton, M. A. C. (1923). On the mammals obtained in Darfur by the LynesLowe expedition. Proceedings of the Zoological Society of London, pp. 247-271. Todd, N. B. (1978). An ecological, behavioural genetic model for the domestication of the cat. Carnivore 1, 52-60. Todd, N. B. & Blumenberg, B. (1978). Mutant allele frequencies and genetic distance relationships in domestic cat populations of lower Egypt and the eastern Mediterranean. Theoretical
and Applied Genetics 52, 257-262.
of Archaeological
Science 1982, 9, 377-379
On the Probable Genotype of Domestic Cats in Ancient Egypt Bennett Blumenberg” Genetic and archaeological evidence suggests that the majority of mummified ritual cats (Felis Kbyca subsp.) in ancient Egypt carried the t + allele and exhibited the striped tabby phenotype. A few of these ritual cats, however, appear to be F. chum subsp., most likely carried the T” allele and were characterized by the Abyssinian phenotype. Severaltypes of evidence suggest that the T”allele first became fixed in a Felis sp. population in either southeast Asia or northeast Africa. A number of Felis subspecies appear to be plausible candidates for the population of T” origin. Keywords: ABYSSINIAN, ALLELE, ANCIENT EGYPT, CHINA, FELZS CATUS, FELZS CHAUS, FELZS LZBYCA OCREATA, GENOTYPE, ALLELE,
PHENOTYPE, POPULATION
GENETICS, STRIPED TABBY, THAILAND.
The paper by Armitage & Clutton-Brock (1981) is a most valuable contribution with respect to (I) the anatomical and histological data established; (2) the elucidation of mummification techniques; and (3) the probable role that domestic cats played in the ritual life of ancient Egyptians. Additional observations that may be of interest can be offered concerning the genotype of these cats. On p. 192, the authors report that microscopic examination established the agouti character of the cat hair. They conclude from this observation that the overall phenotype of these Egyptian cats would have been either striped tabby or Abyssinian. Recent genetic research has established that these two phenotypes are mutually exclusive in Felis sp., for they are controlled by different alleles of the tabby gene (Robinson, 1977). There are three well known forms (alleles) of this gene and they produce the phenotypes known as Abyssinian, striped tabby and blotched tabby. The dominance relationships between these alleles are T” > t f > tb (Robinson, 1977). The t + allele produces the phenotype known as striped tabby that is characterized by vertical curving stripes, short bars or spots on the sides of the body. The head, legs and tail are also distinctively striped; see figure 1 in Robinson (1977). This pattern is believed to represent the ancestral or “wild type” phenotype that is diagnostic of the population (s?) representing the reservoir(s) from which the first domestic groups were drawn (Todd, 1978). Early domestication certainly occurred in Egypt (Armitage & Clutton-Brock, 1981; Baldwin, 1975) and quite possibly in the Indus Valley as well (Ahmad et al., 1980). The Ta allele produces the Abyssinian phenotype in which the striped pattern may be restricted to the head, although frequently found on the legs and tail; see figure 3 in Robinson (1977). While the name “Abyssinian” suggests an origin in northeast Africa,
0305~4403/82/040377+03
“Faculty of Sciences, Lesley College, 29 Everett St, Cambridge, Mass. 02238, U.S.A. 377 $03.00/O 0 1982 Academic Press Inc. (London) Limited
378
B. BLUMENBERG
there is no evidence that this mutant was common in ancient Egypt. Egyptian cat populations now exhibit a genetic profile that is best described as provincial European and the Abyssinian allele is extremely rare or nonexistent (Todd & Blumenberg, 1978). The highest known frequencies of the T” allele ( > 10 %) are found in the isolated Thai city of Mai Hong Son, Hong Kong, Macau and Singapore (Fagen & Meeswat, 1981; Searle, 1959). The likelihood that the Abyssinian allele was present at very low frequencies in the gene pool of early domestic cats in Egypt supports the observation that F. chaus represents very few of those ritual cats that were eventually mummified (Armitage & Clutton-Brock, 1981; Morrison-Scott, 1952). These few F. chaus are significant, however, when hypotheses concerning the origin of the Ta allele are considered (see below). This species is characterized by a phenotype that is essentially devoid of body pattern (Pocock, 1951). F. libyca libyca, which frequently exhibits a well defined tabby pattern (Pocock, 1951, pp. 61-65), appears to represent those populations that quickly came to dominate the domestication process, be this phenotype given distinct species status (Pocock, 1951), believed conspecific with F. silvestris (Todd, 1978) or assigned subspecies rank as F. silvestris libyca (Armitage & Clutton-Brock, 1981); see figures 2 and 3 in Armitage & Clutton-Brock (198 1). Two hypotheses may be offered concerning plausible geographic localities in which the Ta allele first arose and became fixed in Felis sp. populations. Hypothesis 1 suggests a southeast or east Asian origin for this mutant allele and is supported by the biogeographic genetic evidence cited above. F. chaus jiilvidina is found in Burma, Thailand and Vietnam and is characterized by a phenotype virtually devoid of body pattern (Pocock, 1951, pp. 175-176). Pocock (1951, pp. 13-14) describes several feral domestic cat (F. catus) specimens from Burma whose body patterns were faint and obscure. Gray (1972, p. 36) reports that interspecific matings between F. chaus (subspecies not identified) and F. cam produced fertile F, hybrids; such could be the mechanism by which the Ta allele entered the F. catus gene pool. A less attractive variation of this hypothesis could propose that F. bieti bieti, whose “body stripes are sometimes obliterated” (Pocock, 195 1, p. 134) and whose range is the western Chinese provinces of Szechwan and Kansu (now Xinjiang and Xizang), provided the populations of origin for the T” allele. Hypothesis 2 develops from a consideration of the evidence available from Egyptian archaeozoology (Armitage & Clutton-Brock, 1981; Baldwin, 1975; Morrison-Scott, 1952) that points to a northeast African origin for the T” allele. All of these authors agree that a few of the Egyptian ritual cats that were mummified appear to be F. chaus. F. chaus nilotica of Egypt represents one possible source for the T” allele (Flower, 1932, p. 390; Gray, 1972; Pocock, 1951, p. 159). However, it is not necessary to postulate F. chaus as the species in which the T” allele first arose. F. libyca ocreata is essentially devoid of body pattern and has long been known from the Ethiopian region (Allen, 1954, p. 238; Pocock, 1944, pp. 67-68, 1951, pp. 69-72; Thomas & Hinton, 1923, p. 253). Pocock (1907a, b) suggested that the domestic cat (i.e. F. catus) Abyssinian phenotype represents a genetic heritage acquired from F. ocreata (i.e. F. libyca ocreata). Kratochvil & Kratochvil (1976, p. 205) note that F. Zibyca ocreata represents a reasonable alternative to F. libyca libyca as the ancestor of the first domestic cats. Pocock (1907c) reports upon presumed hybrids between F. catus and F. libyca ocreata. If additional evidence serves to firmly establish hypothesis 2, then the origin of the name “Abyssinian” for this particular phenotype will finally have been clarified. Acknowledgements
The author would like to thank Dr J. Clutton-Brock whose comments brought the Gray (1972) reference to his attention and served to stimulate the formulation of origin hypotheses for the T” allele.
GENOTYPE OF DOMESTIC CATS
379
References Ahmad, M., Blumenbcrg, B. & Chaudhary, M. F. (1980). Mutant allele frequencies and genetic distance in cat populations of Pakistan and Asia. Journal of Heredity 71, 323-330. Allen, G. M. (1954). A checklist of African mammals. Bulletin of the Museum of Comparafive Zoology 83, l-763. Armitage, P. L. & Clutton-Brock, J. (1981). A radiological and histological investigation into the mummification of cats from ancient Egypt. Journal of Archaeological Science 8, 185-196. Baldwin, J. A. (1975). Notes and speculations on the domestication of the cat in Egypt. Anrhropos
70, 428-448.
Fagen, R. & Meeswat, K. (1981). Population genetics and demography of the domestic cat (Felis catus L. : Carnivora Felidae) in Thailand. Natural History Bulletin of the Siam Society 29, 7137-150. Flower, S. S. (1932). Notes on the recent mammals of Egypt, with a list of the species recorded from that Kingdom. Proceedings of the Zoological Society of London, pp. 369-450. Gray, A. P. (1972). Mammalian Hybrids: a Checklist with Bibliography. Farnham Royal: Commonwealth Agricultural Bureaux. Kratochvil, J. & Kratochvil, Z. (1976). The origin of domesticated forms of the genus Felis (Mammalia). Zoologicke’ Listy 25, 193-208. Morrison-Scott, T. C. S. (1952). The mummified cats of Ancient Egypt. Proceedings of the Zoological Society of London 121, 861-867. Pocock, R. I. (1907a). On the characters and names of the two types of domestic cat and the name of the European wild cat. Proceedings of rhe Zoological Society of London, pp. 149-155. Pocock, R. I. (19076). On Manx, Persian, Siamese, Indian and Abyssinian breeds. Proceedings of the Zoological Society of London, pp. 161-165. Pocock, R. I. (1907~). On alleged casesof interbreeding between domestic cats and various wild speciesof the genus Felis. Proceedings of the Zoological Society of London, pp. 165-168. Pocock, R. I. (1944). The races of the North African Wild Cat (Felis Lybica). Proceedings of the Zoological Society of London 114, 65-73. Pocock, R. I. (1951). Catalogue of the Genus Felis. London: British Museum. Robinson, R. (1977). Genetics for Car Breeders, 2nd Edn. New York: Pergamon Press. Searle, A. G. (1959). A study of variation in Singapore cats. Journal of Genetics 56, 1-17. Thomas, 0. & Hinton, M. A. C. (1923). On the mammals obtained in Darfur by the LynesLowe expedition. Proceedings of the Zoological Society of London, pp. 247-271. Todd, N. B. (1978). An ecological, behavioural genetic model for the domestication of the cat. Carnivore 1, 52-60. Todd, N. B. & Blumenberg, B. (1978). Mutant allele frequencies and genetic distance relationships in domestic cat populations of lower Egypt and the eastern Mediterranean. Theoretical
and Applied Genetics 52, 257-262.