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Bowling, A. T., Zimmermann, W., Ryder, O., Penado, C., Peto, S., Chemnick, L., et al. (2003). Genetic variation in Przewalski’s horses, with special focus on the last wild caught mare, 231 Orlitza III. Cytogenet Genome Res, 102(1-4), 226–234.
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Lau, A. N., Peng, L., Goto, H., Chemnick, L., Ryder, O. A., & Makova, K. D. (2009). Horse Domestication and Conservation Genetics of Przewalski's Horse Inferred from Sex Chromosomal and Autosomal Sequences. Mol Biol Evol, 26(1), 199–208.
Abstract: Despite their ability to interbreed and produce fertile offspring, there is continued disagreement about the genetic relationship of the domestic horse (Equus caballus) to its endangered wild relative, Przewalski's horse (Equus przewalskii). Analyses have differed as to whether or not Przewalski's horse is placed phylogenetically as a separate sister group to domestic horses. Because Przewalski's horse and domestic horse are so closely related, genetic data can also be used to infer domestication-specific differences between the two. To investigate the genetic relationship of Przewalski's horse to the domestic horse and to address whether evolution of the domestic horse is driven by males or females, five homologous introns (a total of [~]3 kb) were sequenced on the X and Y chromosomes in two Przewalski's horses and three breeds of domestic horses: Arabian horse, Mongolian domestic horse, and Dartmoor pony. Five autosomal introns (a total of [~]6 kb) were sequenced for these horses as well. The sequences of sex chromosomal and autosomal introns were used to determine nucleotide diversity and the forces driving evolution in these species. As a result, X chromosomal and autosomal data do not place Przewalski's horses in a separate clade within phylogenetic trees for horses, suggesting a close relationship between domestic and Przewalski's horses. It was also found that there was a lack of nucleotide diversity on the Y chromosome and higher nucleotide diversity than expected on the X chromosome in domestic horses as compared with the Y chromosome and autosomes. This supports the hypothesis that very few male horses along with numerous female horses founded the various domestic horse breeds. Patterns of nucleotide diversity among different types of chromosomes were distinct for Przewalski's in contrast to domestic horses, supporting unique evolutionary histories of the two species.
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Oakenfull, E. A., Lim, H., & Ryder, O. (2000). A survey of equid mitochondrial DNA: Implications for the evolution, genetic diversity and conservation of Equus. Conservat Genet, 1(4), 341–355.
Abstract: The evolution, taxonomy and conservation of the genus Equuswere investigated by examining the mitochondrial DNA sequences of thecontrol region and 12S rRNA gene. The phylogenetic analysis of thesesequences provides further evidence that the deepest node in thephylogeny of the extant species is a divergence between twolineages; one leading to the ancestor of modern horses (E.ferus, domestic and przewalskii) and the other to thezebra and ass ancestor, with the later speciation events of the zebrasand asses occurring either as one or more rapid radiations, or withextensive secondary contact after speciation. Examination of the geneticdiversity within species suggested that two of the E. hemionussubspecies (E. h. onager and E. h. kulan) onlyrecently diverged, and perhaps, are insufficiently different to beclassified as separate subspecies. The genetic divergence betweendomestic and wild forms of E. ferus (horse) and E.africanus (African ass) was no greater than expected within anequid species. In E. burchelli (plains zebra) there was anindication of mtDNA divergence between populations increasing withdistance. The implications of these results for equid conservation arediscussed and recommendations are made for conservation action.
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Oakenfull, E. A., & Ryder, O. A. (1998). Mitochondrial control region and 12S rRNA variation in Przewalski's horse (Equus przewalskii). Anim Genet, 29(6), 456–459.
Abstract: Variation in the control region and the 12S rRNA gene of all surviving mitochondrial lineages of Przewalski's horse was investigated. Variation is low despite the present day population being descended from 13 individuals probably representing animals from three different regions of its range. Phylogenetic comparison of these sequences, with sequences for the domestic horse, does not resolve the ancestral status of either horse.
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Ryder, O. A., & Wedemeyer, E. A. (1982). A cooperative breeding programme for the mongolian wild horse Equus Przewalski in the United States. Biol. Cons., 22, 259–271.
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Ryder, O. A. (1988). Przewalski's horse – putting the wild horse back in the wild. Oryx, 22, 154–157.
Abstract: The Asian wild horse, or Przewalski's horse (Equus przewalskii), is believed to currently survive only in captivity. There are more than 660 individuals in over 70 zoological collections, and animals are available for reintroduction. The Przewalski's horse had been bred in captivity for 12 generations, and inbreeding has occurred. Genetic variability has been lost, and released animals will require acclimatization on or near release sites. In China, a program is under way for acclimatization, breeding, and release of wild horses, and plans are being made for a similar program in Mongolia. (LCA)
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Ryder, O. A., & Massena, R. (1988). A case of male infanticide in Equus przewalskii. Appl. Anim. Behav. Sci., 21(1-2), 187–190.
Abstract: Following the introduction of a new stallion to a band of E. przewalskii mares two births, both of male foals, resulted in foal death due to injuries sustained in the first day of life. Neither foal was sired by the new herd stallion. The second foal death was the results of an observed attack on the newborn male and is described here. Subsequently births in the same enclosure and, in one instance, to the same mare whose previous foal was killed, were of foals sired by the new stallion and were uneventful, with 3 male foals surviving to date.
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