TY  - THES
A1  - Wutke, Saskia
T1  - Tracing Changes in Space and Time
BT  - Paternal Diversity and Phenotypic Traits during Horse Domestication
N2  - The horse is a fascinating animal symbolizing power, beauty, strength and grace. Among all the animal species domesticated the horse had the largest impact on the course of human history due to its importance for warfare and transportation. Studying the process of horse domestication contributes to the knowledge about the history of horses and even of our own species.
Research based on molecular methods has increasingly focused on the genetic basis of horse domestication. Mitochondrial DNA (mtDNA) analyses of modern and ancient horses detected immense maternal diversity, probably due to many mares that contributed to the domestic population. However, mtDNA does not provide an informative phylogeographic structure. In contrast, Y chromosome analyses displayed almost complete uniformity in modern stallions but relatively high diversity in a few ancient horses. Further molecular markers that seem to be well suited to infer the domestication history of horses or genetic and phenotypic changes during this process are loci associated with phenotypic traits.
This doctoral thesis consists of three different parts for which I analyzed various single nucleotide polymorphisms (SNPs) associated with coat color, locomotion or Y chromosomal variation of horses. These SNPs were genotyped in 350 ancient horses from the Chalcolithic (5,000 BC) to the Middle Ages (11th century). The distribution of the samples ranges from China to the Iberian Peninsula and Iceland. By applying multiplexed next-generation sequencing (NGS) I sequenced short amplicons covering the relevant positions: i) eight coat-color-associated mutations in six genes to deduce the coat color phenotype; ii) the so-called ’Gait-keeper’ SNP in the DMRT3 gene to screen for the ability to amble; iii) 16 SNPs previously detected in ancient horses to infer the corresponding haplotype. Based on these data I investigated the occurrence and frequencies of alleles underlying the respective phenotypes as well as Y chromosome haplotypes at different times and regions. Also, selection coefficients for several Y chromosome lineages or phenotypes were estimated.
Concerning coat color differences in ancient horses my work constitutes the most comprehensive study to date. I detected an increase of chestnut horses in the Middle Ages as well as differential selection for spotted and solid phenotypes over time which reflects changing human preferences.
With regard to ambling horses, the corresponding allele was present in medieval English and Icelandic horses. Based on these results I argue that Norse settlers, who frequently invaded parts of Britain, brought ambling individuals to Iceland from the British Isles which can be regarded the origin of this trait. Moreover, these settlers appear to have selected for ambling in Icelandic horses.
Relating to the third trait, the paternal diversity, these findings represent the largest ancient dataset of Y chromosome variation in non-humans. I proved the existence of several Y chromosome haplotypes in early domestic horses. The decline of Y chromosome variation coincides with the movement of nomadic peoples from the Eurasian steppes and later with different breeding practices in the Roman period.
In conclusion, positive selection was estimated for several phenotypes/lineages
in different regions or times which indicates that these were preferred by humans. Furthermore, I could successfully infer the distribution and dispersal of horses in association with human movements and actions. Thereby, a better understanding of the influence of people on the changing appearance and genetic diversity of domestic horses could be gained. My results also emphasize the close relationship of ancient genetics and archeology or history and that only in combination well-founded conclusions can be reached.
KW  - ancient DNA
KW  - domestication
KW  - horse
KW  - equus caballus
KW  - locomotion
KW  - Y chromosome
KW  - coat colour
Y1  - 2016
ER  - 
TY  - GEN
A1  - Wutke, Saskia
A1  - Andersson, Leif
A1  - Benecke, Norbert
A1  - Sandoval-Castellanos, Edson
A1  - Gonzalez, Javier
A1  - Hallsson, Jon Hallsteinn
A1  - Lougas, Lembi
A1  - Magnell, Ola
A1  - Morales-Muniz, Arturo
A1  - Orlando, Ludovic
A1  - Palsdottir, Albina Hulda
A1  - Reissmann, Monika
A1  - Munoz-Rodriguez, Mariana B.
A1  - Ruttkay, Matej
A1  - Trinks, Alexandra
A1  - Hofreiter, Michael
A1  - Ludwig, Arne
T1  - The origin of ambling horses
T2  - Current biology
N2  - Horseback riding is the most fundamental use of domestic horses and has had a huge influence on the development of human societies for millennia. Over time, riding techniques and the style of riding improved. Therefore, horses with the ability to perform comfortable gaits (e.g. ambling or pacing), so-called ‘gaited’ horses, have been highly valued by humans, especially for long distance travel. Recently, the causative mutation for gaitedness in horses has been linked to a substitution causing a premature stop codon in the DMRT3 gene (DMRT3_Ser301STOP) [1]. In mice, Dmrt3 is expressed in spinal cord interneurons and plays an important role in the development of limb movement coordination [1]. Genotyping the position in 4396 modern horses from 141 breeds revealed that nowadays the mutated allele is distributed worldwide with an especially high frequency in gaited horses and breeds used for harness racing [2]. Here, we examine historic horse remains for the DMRT3 SNP, tracking the origin of gaitedness to Medieval England between 850 and 900 AD. The presence of the corresponding allele in Icelandic horses (9th–11th century) strongly suggests that ambling horses were brought from the British Isles to Iceland by Norse people. Considering the high frequency of the ambling allele in early Icelandic horses, we believe that Norse settlers selected for this comfortable mode of horse riding soon after arrival. The absence of the allele in samples from continental Europe (including Scandinavia) at this time implies that ambling horses may have spread from Iceland and maybe also the British Isles across the continent at a later date.
Y1  - 2016
U6  - https://doi.org/10.1016/j.cub.2016.07.001
SN  - 0960-9822
SN  - 1879-0445
VL  - 26
SP  - R697
EP  - R699
PB  - Cell Press
CY  - Cambridge
ER  - 
TY  - JOUR
A1  - Wutke, Saskia
A1  - Benecke, Norbert
A1  - Sandoval-Castellanos, Edson
A1  - Döhle, Hans-Jürgen
A1  - Friederich, Susanne
A1  - Gonzalez Soto, Javier Esteban
A1  - Hallsson, Jon Hallsteinn
A1  - Hofreiter, Michael
A1  - Lougas, Lembi
A1  - Magnell, Ola
A1  - Morales-Muniz, Arturo
A1  - Orlando, Ludovic
A1  - Palsdottir, Albina Hulda
A1  - Reissmann, Monika
A1  - Ruttkay, Matej
A1  - Trinks, Alexandra
A1  - Ludwig, Arne
T1  - Spotted phenotypes in horses lost attractiveness in the Middle Ages
JF  - Scientific reports
N2  - Horses have been valued for their diversity of coat colour since prehistoric times; this is especially the case since their domestication in the Caspian steppe in similar to 3,500 BC. Although we can assume that human preferences were not constant, we have only anecdotal information about how domestic horses were influenced by humans. Our results from genotype analyses show a significant increase in spotted coats in early domestic horses (Copper Age to Iron Age). In contrast, medieval horses carried significantly fewer alleles for these phenotypes, whereas solid phenotypes (i.e., chestnut) became dominant. This shift may have been supported because of (i) pleiotropic disadvantages, (ii) a reduced need to separate domestic horses from their wild counterparts, (iii) a lower religious prestige, or (iv) novel developments in weaponry. These scenarios may have acted alone or in combination. However, the dominance of chestnut is a remarkable feature of the medieval horse population.
Y1  - 2016
U6  - https://doi.org/10.1038/srep38548
SN  - 2045-2322
VL  - 6
PB  - Nature Publ. Group
CY  - London
ER  - 
TY  - JOUR
A1  - Patel, Riddhi P.
A1  - Wutke, Saskia
A1  - Lenz, Dorina
A1  - Mukherjee, Shomita
A1  - Ramakrishnan, Uma
A1  - Veron, Geraldine
A1  - Fickel, Jörns
A1  - Wilting, Andreas
A1  - Förster, Daniel W.
T1  - Genetic Structure and Phylogeography of the Leopard Cat (Prionailurus bengalensis) Inferred from Mitochondrial Genomes
JF  - Journal of Heredity
N2  - The Leopard cat Prionailurus bengalensis is a habitat generalist that is widely distributed across Southeast Asia. Based on morphological traits, this species has been subdivided into 12 subspecies. Thus far, there have been few molecular studies investigating intraspecific variation, and those had been limited in geographic scope. For this reason, we aimed to study the genetic structure and evolutionary history of this species across its very large distribution range in Asia. We employed both PCR-based (short mtDNA fragments, 94 samples) and high throughput sequencing based methods (whole mitochondrial genomes, 52 samples) on archival, noninvasively collected and fresh samples to investigate the distribution of intraspecific genetic variation. Our comprehensive sampling coupled with the improved resolution of a mitochondrial genome analyses provided strong support for a deep split between Mainland and Sundaic Leopard cats. Although we identified multiple haplogroups within the species’ distribution, we found no matrilineal evidence for the distinction of 12 subspecies. In the context of Leopard cat biogeography, we cautiously recommend a revision of the Prionailurus bengalensis subspecific taxonomy: namely, a reduction to 4 subspecies (2 mainland and 2 Sundaic forms).
KW  - habitat generalist
KW  - hybrid capture
KW  - Leopard cat
KW  - mitogenome
KW  - mtDNA
KW  - Southeast Asia
Y1  - 2017
U6  - https://doi.org/10.1093/jhered/esx017
SN  - 0022-1503
SN  - 1465-7333
VL  - 108
IS  - 4
SP  - 349
EP  - 360
PB  - Oxford Univ. Press
CY  - Cary
ER  - 
TY  - JOUR
A1  - Wutke, Saskia
A1  - Sandoval-Castellanos, Edson
A1  - Benecke, Norbert
A1  - Döhle, Hans-Jürgen
A1  - Friederich, Susanne
A1  - Gonzalez, Javier
A1  - Hofreiter, Michael
A1  - Lougas, Lembi
A1  - Magnell, Ola
A1  - Malaspinas, Anna-Sapfo
A1  - Morales-Muniz, Arturo
A1  - Orlando, Ludovic
A1  - Reissmann, Monika
A1  - Trinks, Alexandra
A1  - Ludwig, Arne
T1  - Decline of genetic diversity in ancient domestic stallions in Europe
JF  - Science Advances
N2  - Present-day domestic horses are immensely diverse in their maternally inherited mitochondrial DNA, yet they show very little variation on their paternally inherited Y chromosome. Although it has recently been shown that Y chromosomal diversity in domestic horses was higher at least until the Iron Age, when and why this diversity disappeared remain controversial questions. We genotyped 16 recently discovered Y chromosomal single-nucleotide polymorphisms in 96 ancient Eurasian stallions spanning the early domestication stages (Copper and Bronze Age) to the Middle Ages. Using this Y chromosomal time series, which covers nearly the entire history of horse domestication, we reveal how Y chromosomal diversity changed over time. Our results also show that the lack of multiple stallion lineages in the extant domestic population is caused by neither a founder effect nor random demographic effects but instead is the result of artificial selection-initially during the Iron Age by nomadic people from the Eurasian steppes and later during the Roman period. Moreover, the modern domestic haplotype probably derived from another, already advantageous, haplotype, most likely after the beginning of the domestication. In line with recent findings indicating that the Przewalski and domestic horse lineages remained connected by gene flow after they diverged about 45,000 years ago, we present evidence for Y chromosomal introgression of Przewalski horses into the gene pool of European domestic horses at least until medieval times.
Y1  - 2018
U6  - https://doi.org/10.1126/sciadv.aap9691
SN  - 2375-2548
VL  - 4
IS  - 4
PB  - American Assoc. for the Advancement of Science
CY  - Washington
ER  -