The Origin of African Sheep: Archaeological and Genetic Perspectives

Abstract

Indigenous African sheep genetic resources have been classified into two main groups, fat-tailed and thin-tailed sheep. The fat-tailed sheep are the most widely distributed, being found in a large part of North Africa (from Egypt to Algeria) and in Eastern and Southern Africa (from Eritrea to South Africa). The thin-tailed sheep are present mainly in Morocco, Sudan and in West Africa. African sheep were domesticated outside Africa. They share a common ancestry with European and Asian sheep. Archaeological information supports separate introductions and dispersion histories for the African thin-tailed and fat-tailed sheep. The first sheep entered Africa via the Isthmus of Suez and/or the southern Sinai Peninsula, between 7500 and 7000 BP. They were likely of the thin-tailed type. Fat-tailed sheep entered Africa through its northeastern part and the Horn of Africa. Mitochondrial DNA analysis supports a common maternal ancestral origin for all African sheep, while autosomal and Y chromosome DNA analysis indicates a distinct genetic history for African thin-tailed and sub-Saharan fat-tailed sheep. The main ancestral population of southern African fat-tailed sheep likely originated in East Africa. Further work is needed to assess the possible dispersion of sheep from western Africa to the southern African regions.

Full text

REVIEW ARTICLE
The Origin of African Sheep: Archaeological
and Genetic Perspectives
Anne W. T. Muigai &Olivier Hanotte
Published online: 6 March 2013
#Springer Science+Business Media New York 2013
Abstract Indigenous African sheep genetic resources have been classified into two
main groups, fat-tailed and thin-tailed sheep. The fat-tailed sheep are the most widely
distributed, being found in a large part of North Africa (from Egypt to Algeria) and in
Eastern and Southern Africa (from Eritrea to South Africa). The thin-tailed sheep are
present mainly in Morocco, Sudan and in West Africa. African sheep were domes-
ticated outside Africa. They share a common ancestry with European and Asian
sheep. Archaeological information supports separate introductions and dispersion
histories for the African thin-tailed and fat-tailed sheep. The first sheep entered
Africa via the Isthmus of Suez and/or the southern Sinai Peninsula, between 7500
and 7000 BP. They were likely of the thin-tailed type. Fat-tailed sheep entered Africa
through its northeastern part and the Horn of Africa. Mitochondrial DNA analysis
supports a common maternal ancestral origin for all African sheep, while autosomal
and Ychromosome DNA analysis indicates a distinct genetic history for African thin-
tailed and sub-Saharan fat-tailed sheep. The main ancestral population of southern
African fat-tailed sheep likely originated in East Africa. Further work is needed to
assess the possible dispersion of sheep from western Africa to the southern African
regions.
Résumé Les ressources génétiques africaines du mouton sont classifiées en deux
groupes principaux, les moutons à queue grasse et les moutons à queue fine. Les
moutons à queue grasse ont la distribution géographique la plus étendue, se
rencontrant dans une grande partie de l’Afrique du Nord (depuis l’Égypte jusqu’à
Afr Archaeol Rev (2013) 30:39–50
DOI 10.1007/s10437-013-9129-0
A. W. T. Muigai (*)
Jomo Kenyatta University of Agriculture and Technology, P.O. Box 62000, Nairobi 00200, Kenya
e-mail: awmuigai@yahoo.co.uk
O. Hanotte
School of Biology, Centre for Genetics and Genomics, University of Nottingham, University Park,
Nottingham NG7 2RD, UK

l’Algérie), et dans les régions les plus à l’est et au sud de l’Afrique (de l’Érythrée à
l’Afrique du Sud). Les moutons à queue fine sont présents principalement au Maroc,
au Soudan et en Afrique de l’Ouest. Les moutons africaines ont été domestiqués en
dehors de l’Afrique. Ils partagent un ancêtre commun avec les moutons asiatiques et
européens. Les informations archéologiques supportent une introduction distincte et
une histoire de dispersion en partie séparée pour les moutons à queue grasse et à
queue fine sur le continent. Les premiers moutons sont arrivés en Afrique par
l’Isthme de Suez et/ou le sud de la Péninsule du Sinaï, entre 7500 et 7000 BP. Ils
étaient probablement du type mouton à queue fine. Les moutons à queue grasse
entrèrent en Afrique par le nord-est du continent et la corne de l’Afrique. Les résultats
de l’ADN mitochondrial supportent une origine ancestrale maternelle commune pour
tous les moutons africaines, tandis que les données des autosomes et du chromosome
Yindiquent une histoire génétique distincte pour les moutons à queue grasse et à
queue fine. La population ancestrale principale des moutons à queue grasse du sud du
continent est probablement originaire de l’est de l’Afrique. Des études supplémentaires
sont nécessaires pour attester la dispersion possible du mouton de l’Afrique de l’Ouest
jusqu’au sud du continent.
Keywords Africa .Indigenous sheep .Origin .Migration .Genetic relationships
Introduction
The present-day, sub-Saharan African domestic sheep (Ovis aries) genetic resources
are extensive, with an estimated 170 breeds (Kemp et al.2007). Eighty percent of the
sheep are classified as indigenous populations maintained under traditional farming
systems (Kosgey et al.2008). They are divided into two main groups with a largely
non-overlapping distribution: thin-tailed and fat-tailed (including fat-rump) sheep
(Wilson 1991; Rege et al.1996).
The thin-tailed sheep are found mainly in North Africa from Morocco to Egypt, in
Sudan and in West Africa from Sénégal to Nigeria. There are two main sub-types, the
African long-legged, found mainly in the arid and semi-arid zones of the Sahel and
Western Africa (Fig. 1a), and the tropical dwarf sheep (Djallonké sheep) found
mainly in the sub-humid and humid zones of West and Central Africa (Fig. 1b).
They are considered to be the most ancient type of sheep on the continent and their
convex nose, pendulous ears and exceptionally long legs closely resemble the sheep
of ancient Egypt (Ryder 1983,1984; Wilson 1991). The fat-tailed sheep are the most
widely distributed sheep on the continent today. They are found in northern Africa
from Egypt to Algeria, and in the eastern and the southern regions of the continent
from Eritrea to South Africa. These sheep are further sub-divided into two sub-types,
the fat-tailed hair (Fig.1c) and fat-tailed wool (Fig. 1d) sheep, depending on types of
coat (Rege et al.1996).
Like other domestic sheep, African sheep are thought to descend from the Asiatic
mouflon (Ovis orientalis) (Bruford and Townsend 2006; Meadows et al.2011). They
were most likely domesticated outside the continent, as wild Ovis species are not
native to Africa. An African centre of domestication has been proposed (Muzzolini
1987,1993,2000), but it is not currently supported by any archaeological or genetic
40 Afr Archaeol Rev (2013) 30:39–50

data (Clutton-Brock 1999; MacDonald and MacDonald 2000; Kijas et al.2012).
Domestication of sheep is thought to have begun in the central Fertile Crescent,
especially the upper parts of the Tigris and Euphrates rivers (Lev-Yadun et al.2000;
Zeder 2011).
Domestication of wild animals is no longer viewed as an event or an invention but
rather a co-evolutionary process (e.g., Gifford-Gonzalez and Hanotte 2011). There
are now emerging archaeological and genetic data showing that domestication of
ruminant species began as low-intensity management of wild animals, followed by
more intensive herd management and finally full domestication (Zeder 2011). The
domestication process could have therefore begun much early than the manifestation
of archaeologically detectable morphological changes in such managed populations
(Zeder 2011). The sheep remains from Körtik Tepe in present-day Turkey (c.
10,900 cal BP) have been interpreted as being in a transitional strategy between
game management and herd management (Arbuckle and Özkaya 2006), and a similar
interpretation for the sheep remains from Zawi Chemi Shanidar in the north-western
Zagros, of the late Epipaleolithic (c. 12,000–11,500 cal BP), has now been proposed
(Zeder 2011). Caprine remains from Nevali Çori (Turkey) give an indication that the
transition from hunting to herding was complete by 10,500 cal BP, with sheep being
an earlier focus of herd management than goats (Zeder 2011).
The wild ancestor of O.aries is thin-tailed, and accordingly the first domesticated
sheep were most likely thin-tailed. The first depiction of fat-tail sheep, on an Uruk II
stone vessel dated from 3000 BC, is several thousand years after the first evidence of
sheep domestication (Ryder 1983). Thin-tailed sheep were the first to move outside
their centre of domestication, with groups migrating northward and along the
Fig. 1 The sheep of Africa. aThin-tailed African long-legged—Uda; bThin-tailed Tropical Dwarf—
Djallonké; cFat-tailed hair—Red Maasai; dFat-tailed wool—Menz (©DAGRIS, 2007)
Afr Archaeol Rev (2013) 30:39–50 41

Mediterranean basin, giving rise to present-day European thin-tailed breeds as well as
the feral populations of the Mediterranean islands (Epstein 1971; Pereira et al.2006;
Chessa et al.2009). Interestingly, today fat-tailed sheep are prevalent in the Fertile
Crescent, at the geographic region of sheep domestication, while thin-tailed sheep are
most commonly found on the periphery of the geographic distribution of modern
sheep (Ryder 1984). The origins of fat-tailed sheep remain uncertain but, given their
modern geographic range and the absence of putative fat-tailed wild ancestors, it is
likely that the fat-tailed phenotype was selected for in domestic populations of
originally thin-tailed sheep, possibly in response to extreme climatic conditions
and/or human preferences (Epstein 1971; Moradi et al.2012).
Here, we review the available archaeological and molecular genetic information on
the history and diversity of indigenous African sheep in the broader context of our
current knowledge on sheep domestication and worldwide dispersion, with the aim to
guide further research on the African history of this domesticated species.
The Origin and Migration of Domesticated Sheep into Africa: Archaeological
Evidence
Evidence of sheep on the African continent is much more recent than the Southwest
Asian occurrences at the domestication centres. Faunal remains, as well as ancient
paintings and rock carvings across the continent,are providing insight on the geographic
entry points and subsequent history of sheep husbandry on the continent. The earliest
African domestic sheep remains, dated between 7500and 7000 BP, derivefrom the Nile
Delta, the eastern Sahara and the Red Sea Hills (Gautier 1987;Close1992; Vermeersch
et al.1996). These sheep likely originated from the Sinai Peninsula, where sheep
remains have been dated c. 7000 BP (Oren 1979). They entered into the African
continent, perhaps in response to droughts and unstable climates in the Levant. Sheep
remains have been positively identified in the Sodmein Cave in the Red Sea Hills with
radiocarbon dates 7100–7000 BP (Vermeersch et al.1994; Vermeersch et al.1996). The
overland dispersal route of sheep was northward to Libya (6500–6800 BP), southwards
to that central Nile valley (6000 BP) and westwards to that central Sahara (6000 BP)
reaching West Africa by 3700 BP (Close 1992;Smith1992; Breunig et al.1996;Holl
1998;Hassan2000; MacDonald and MacDonald 2000). Mediterranean maritime trad-
ing routes may have facilitated the dispersal of sheep along the northern African
coastline (Fig. 2; Ryder 1984; Pereira et al.2006).
The depiction of animal forms in ancient rock engravings and paintings provides
information on the phenotypes and appearance of these first African sheep. The
Tassili rock paintings show flocks of horned sheep with hair coats and thin tails.
Numerous representations of sheep are found during the Early Dynastic Period of
Egypt (5111–4624 BP). One of the earliest of these pictures is a pre-dynastic Tjehnu
palette, dated 5000 BP, in which domesticated sheep with corkscrew horns and long
thin tails are represented together with domesticated cattle and asses (Gardiner 1961).
Generally, the paintings of sheep from ancient Egyptian tombs from the Early
Dynastic Period of Egypt depict sheep with a straight head profile and Roman nose,
spiral horns, long thin legs, a long thin tail reaching the hocks and often black, white
or pied coat (Epstein 1971;ChaixandGrant1987;Marshall2000). Their phenotypic
42 Afr Archaeol Rev (2013) 30:39–50

resemblances with the corkscrew-horned sheep with long legs, lop ears and pied colours
found presently in West Africa suggest that the latter may be the descendants of these
ancient Egyptian sheep, for example the Uda breed of West Africa, characterised by a
striking two-colour coat pattern, long thin tail and long legs (Fig. 1a).
The sheep inhabiting northern Africa are today all fat-tailed. We cannot fully exclude
the possibility that thisphenotype might have emerged independently from the fat-tailed
sheep of the Fertile Crescent, but we think it is unlikely in light of the continuous
geographic distribution of these fat-tailed populations with their Asian counterparts. The
distribution instead suggests that a second major wave of sheep entered North Africa and
subsequently replaced and/or introgressed with the local thin-tailed sheep. Accordingly,
fat-tailed sheep are depicted more recently in Egypt compared to thin-tailed sheep,
during the Middle Kingdom, 3002–2644 BP (Clutton-Brock 1993). Fat-tailed sheep are
not found in West Africa. There is, to the best of our knowledge, no evidence of fat-
tailed sheep in western Saharan rock paintings and/or engravings.
D
?
Fig. 2 Possible routes of introduction and spread of sheep pastoralism into Africa. Thin- and fat-tailed
sheep likely originate from the same broad geographic zone in the Zagros Mountains of Southwest Asia
(Dregion of domestication). Thin-tailed sheep (light line) were the first to migrate from their centre of
domestication, entering the African continent via the Isthmus of Suez and diffusing overland and through
maritime trade (dark dotted line). Fat-tailed sheep (dark continuous line) entered the continent via the
Isthmus of Suez and from the Arabian Peninsula via the Horn of Africa, migrating southward. Another
movement of sheep from Western Africa to Southern Africa along the Atlantic coastline (light dashed line)
is also possible but needs to be confirmed (see text for details and citations)
Afr Archaeol Rev (2013) 30:39–50 43

The spread of sheep into East Africa is less well documented. By 4500–3500 BP,
sheep are found with cattle in the Lake Turkana basin (Barthelme 1985; MacDonald
and MacDonald 2000). Clark (1976) suggests that these sheep may have diffused
overland from the Nile Valley in response to growing aridity in the north. If so, these
would have been sheep of the thin-tailed type, as fat-tailed sheep were not present in
the Nile Valley at the time. However, there are also increasing indications of direct
trading connections between the Arabian Peninsula and the Eritrean region, including
domestic animals (Marshall 2000; Boivin and Fuller 2009). The first East African
sheep could therefore have been introduced directly from the Arabian Peninsula.
Undated rock paintings from Serkama Cave (Harar Province) in the eastern Ethiopian
highlands (Clark and Williams 1978) portray fat-tailed sheep and humpless cows,
suggesting that fat-tailed sheep were present in East Africa prior the arrival of
humped cattle, for which there is undisputed archaeological evidence dated to around
the mid-first millennium AD (Gifford-Gonzalez and Hanotte 2011). It remains
unknown if the arrival of fat-tailed sheep in East Africa and in the northeastern part
of the continent occurred in the same time period.
Sheep arrived in Southern Africa in approximately the late third millennium BP
(Sealy and Yates 1994; Sealy and Yates 1996), reaching the Western Cape ca. 2000
BP (Smith 2009). Southern African sites yielding sheep remains have been associated
with Later Stone Age and early pastoralist Khoi communities (Plug and Badenhorst
2001) as well as Early Iron Age and/or Early Farming communities (Mitchell 2002).
The oldest evidence of caprines in Southern Africa is from the Later Stone Age (LSA)
site of Leopard Cave, Erongo, Namibia (Pleurdeau et al.2012). It has been dated
from 2270±40 BP. Whether the two teeth found represent sheep or goat remains
unclear, but the positive identification of sheep bones in the overlying layer supports
the presence of sheep rather than goat at the site (Pleurdeau et al.2012). In south-
ernmost Africa, the earliest evidence of sheep husbandry dates from1960±50 BP
from the Blombos site on the Southern Cape coast (Henshilwood 1996). The route
that livestock took to reach Southern Africa is still debated, and several hypotheses
have been proposed including migrations through and along the coastal areas of the
Congo Basin and through tsetse corridors along the highland of the Rift Valley
(Fig. 2; Smith 1998,2009; Gifford-Gonzalez and Hanotte 2011). Also, it remains
debated whether the appearance of caprines in Southern Africa involved immigration
of herders from the north or if it was the result of a cultural diffusion process.
The tail phenotype of the first Southern African sheep is unknown. Undated rock
art depictions of both fat-tailed and thin-tailed sheep have been reported from
Zimbabwe (Goodall 1946 cited in Rocha et al.2011) and Southern African rock art
depicts sheep as fat-tailed with a hair coat (Campbell and Coulson 2001), similar to
present-day East African sheep.
The Origin and Migration of Domesticated Sheep into Africa: Molecular
Genetic Evidence
Domestication and dispersion of livestock are studied through the analysis of mito-
chondrial DNA as well as autosomal and Y-specific DNA markers (Bruford et al.
2003). Mitochondrial DNA analysis initially revealed at least two distinct maternal
44 Afr Archaeol Rev (2013) 30:39–50

lineages (haplogroups A and B) in domestic sheep (Wood and Phua 1996; Hiendleder
et al.1998). Later, three more lineages (haplogroups C, D, E) were found (Guo et al.
2005; Pedrosa et al.2005; Tapio et al.2006; Meadows et al.2007). Today, the
complete mitogenome of these haplogroups has been sequenced, showing that they
all shared a common maternal ancestry distinct from the argali (O.ammon) and urial
(O.vignei), leaving the Asian mouflon (O.orientalis) as the most likely maternal
ancestor of all domestic sheep (Meadows et al.2011). It remains unknown if these
genetically distinct mitochondrial DNA lineages relate to one or several distinct
domestication events. However, the former is perhaps more likely, in light of the
increased awareness that domestication of livestock was likely preceded by a man-
agement phase of wild populations across an extended geographic area (Zeder 2011).
Interestingly, a recent genome-wide analysis of sheep breeds across the world has
revealed high levels of genetic admixture compatible with a single large ancestral
genetic pool for all domestic sheep (Kijas et al.2012).
Compared to European and Asian sheep, the diversity of mitochondrial DNA of
African sheep has been less studied. All the mtDNA data so far indicate that African
sheep share a common maternal ancestry with European and Asian sheep, and that they
likely originated from the same centre(s) of domestication. Bruford and Townsend (2006)
studied the mtDNA D-loop diversity of eight East and Southern African fat-tailed sheep
populations and identified haplogroup A only. However, Horsburgh and Rhines (2010),
partly sequencing the mtDNA control region of 22 sheep bone samples from Die Kelders
Cave in South Africa, found that all samples belonged to haplogroup B. The dating of
these samples remains uncertain, but in no case are they older than 2000 years. Recently,
Gornas et al.(2011) characterised the mitochondrial DNA of 231 thin-tailed Sudanese
sheep, identifying haplogroup B in 90 % of the animals and haplogroup A in the
remaining 10 %. Mitochondrial haplotype DNA diversity of Nigerian sheep has been
reported (Agaviezor et al.2012a), but unfortunately no information was provided regard-
ing their haplogroup memberships. All genetic evidence so far therefore indicates a
common maternal ancestry for African sheep across the continent.
The first molecular study to show genetic differentiation between West African thin-
tailed sheep and the fat-tailed sheep of East and Southern Africa was the study by
Muigai (2003), using autosomal microsatellite markers. This represents the most ex-
tended study thus far, both in terms of geographic coverage and the number of breeds
sampled, of the autosomal genetic diversity of African sheep. Twenty-four sheep
populations, 16 indigenous to the African continent and eight that reference populations
from outside the continent, were studied. The indigenous African populations included
seven fat-tailed breeds from eastern Africa (Red Maasai, Afar, Horro, Menz, Tekur,
Wello and Newala), four fat-tailed breeds from southern Africa (Damara, Sabi, Tswana
and Pedi), one fat-tailed breed from North Africa (Ossimi) and three thin-tailed sheep
breeds from West Africa (Maure, Uda, Djallonké sampled in Sénégal and Djallonké
sampled in Nigeria). The non-African populations included two Middle Eastern
populations: Awassi (fat-tailed sheep sampled in Syria) and Mahli (fat-tailed sheep
sampled in the United Arab Emirates). Three European thin-tailed populations
(Dorset, Swaledale, North Ronaldsay), two Asian fat-tailed populations (Lanzhou,
Karakul) and one Indian thin-tailed sheep (Deccani) were also studied.
Thirty-five to 40 unrelated individuals from each population were analyzed. Allele
frequencies at the microsatellite loci were calculated and used to determine the
Afr Archaeol Rev (2013) 30:39–50 45

genetic relationships and population structure of the studied populations. The results
for the African and non-African breeds are reported in detail in Muigai (2003).
Interestingly, they show: (1) a clear separation between the fat-tailed African popu-
lation from East and Southern Africa and all the remaining West African sheep and
non-African populations studied; (2) a closer relationship between the North African
fat-tailed, coarse wool Ossimi and the thin-tailed West African hair sheep than with
any East and Southern African fat-tailed sheep; (3) a common autosomal genetic
background for sub-Saharan fat-tailed sheep and the Mahli, the only Arabian
Peninsula fat-tailed sheep sampled; and (4) the closest relations of Southern
African fat-tailed sheep are with the East African fat-tailed sheep populations.
Together, these results indicate that the sub-Saharan African fat-tailed sheep represent
a separate genetic stock at the autosomal level, agreeing with archaeological infor-
mation showing a distinct history for the West African thin-tailed and the fat-tailed
sheep from East and Southern Africa. The study also called for further research to
address the issue of a possible common genetic background between Southern
African fat-tailed and West African thin-tailed sheep.
At the Ychromosome, two studies have looked at polymorphic loci in African
sheep (Meadows et al.2006; Aswani 2007). Using combined information from a
single nucleotide polymorphic and one microsatellite marker (SRYMA18), Meadows
et al.(2006) identified five Ychromosome haplotypes in a total of 42 animals
representative of West, East and Southern African sheep. Aswani (2007) expanded
this study by analyzing 447 thin-tailed and fat-tailed African sheep. He found that the
frequency of Ychromosome haplotypes differed between West Africa on the one
hand and East and Southern Africa on the other, adding further support to the
archaeological and autosomal genetic information suggesting distinct histories.
It is also worth noting that, in a study using endogenous retroviruses as genetic
markers, Chessa et al. (2009) found the same retroviral signature in “primitive”
European sheep and the Djallonké sheep, a thin-tailed sheep from The Gambia.
This finding needs to be confirmed with the analysis of more African sheep
populations, but it may indicate that African thin-tailed and European primitive sheep
breeds may represent a common modern legacy, of the first dispersion of domestic
sheep from their centres of domestication to Europe and Africa.
Finally, several country- or region-specific microsatellite studies have been now
performed on African sheep (Muigai 2003; Muigai et al. 2009; Agaviezor et al.
2012b;Somaet al.2012). Though restricted in their geographic scope, these studies
indicate close relationships and high levels of admixture among sheep populations
from the same geographic area.
Discussion and Conclusion
Through a combination of archaeological and molecular genetics findings, the history
of African sheep pastoralism is emerging (Fig. 2). Domestic sheep entered the
continent through two main entry points: the northeastern part of Africa and the
Horn of Africa, with the latter a further illustration of the importance of that region in
the history of African livestock pastoralism (Hanotte et al.2002; Boivin and Fuller
2009). However, despite rock painting evidence that fat-tailed sheep might have been
46 Afr Archaeol Rev (2013) 30:39–50

present in the Horn of Africa prior the arrival and dispersion of humped cattle along
the Sahelian belt into West Africa (Clark and Williams 1978), fat-tailed sheep did not
reach the western part of the continent along the same route. Depictions of thin-tailed
and fat-tailed sheep at different periods in Dynastic Egyptian art reveal several waves
of introduction in the northeastern part of the continent. Last, genetic information
indicates a largely distinct separate history for West African thin-tailed sheep from
East or Southern African fat-tailed sheep (Muigai 2003; Aswani 2007).
However, many unknowns remain, particularly the pattern and process of the diffu-
sion of sheep pastoralism across the continent. Did sheep disperse alone or in the
company of other major livestock such as goat and/or cattle? Was it a process of cultural
diffusion and adoption, or rather the result of migratory movements of pastoral com-
munities? Were the first Southern African sheep of the fat-tailed or thin-tailed type? Did
they reach the southern part of the continent through tsetse corridors along the highlands
of the Rift Valley as well as along the coastal areas of the Congo Basin?
New archaeological information and ancient DNA studies may further clarify
these issues. However, so far, we lack a genetic marker for the fat-tailed/thin-tailed
phenotypes, as they do not correlate with the major haplotypes in Africa. Nor are
there diagnostic skeletal criteria to separate the two. These deficiencies will continue
to limit our understanding of the history of sheep on the African continent. The work
of Moradi et al.(2012) is a first step toward the possible identification of such
markers. When available, they will undoubtedly provide new, perhaps unexpected,
light on the history of African sheep.
Acknowledgments The authors wish to thank Diane Gifford-Gonzalez for her very useful comments and
sharp-eyed editing, and the livestock keepers of Africa for preserving the unique and diverse sheep genetic
resources of Africa to present times.
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