Book Review Essays

The Ark of the Covenant in Africa: The Beta Israel (Falasha) in Ethiopia: From Earliest Times to the Twentieth Century. Steven Kaplan.
The Ark of the Covenant in Africa: The Evolution of the Ethiopian Jews: A History of the Beta Israel (Falasha) to 1920. James Quirin.     

The differences among Jewish communities—Maternal and paternal contributions

The haplotypes of Y chromosome (paternally inherited) and mtDNA (maternally inherited) were analyzed in representatives of six Jewish communities (Ashkenazic, North African, Near Eastern, Yemenite, Minor Asian/Balkanian, and Ethiopian). For both elements, the Ethiopian community has a mixture of typically African and typically Caucasian haplotypes and is significantly different from all others. The other communities, whose haplotypes are mostly Caucasian, are more closely related; significant differences that were found among some of them possibly indicate the effects of admixture with neighboring communities of non-Jews. The different contribution of the Y chromosome and mtDNA haplotypes to the significant differences among the communities can be explained by unequal involvement of males and females in the different admixtures. In all communities, except the Ethiopians, the level of diversity () for Y chromosome haplotypes is higher than that for mtDNA haplotypes, suggesting that in each community the people who become parents include more males than females. An opposite proportion (more females than males) is found among the Ethiopians. Correspondence to: U. Ritte     

Mitochondrial DNA variation reveals maternal origins and demographic dynamics of Ethiopian indigenous goats

The Horn of Africa forms one of the two main historical entry points of domestics into the continent and Ethiopia is particularly important in this regard. Through the analysis of mitochondrial DNA (mtDNA) d‐loop region in 309 individuals from 13 populations, we reveal the maternal genetic variation and demographic dynamics of Ethiopian indigenous goats. A total of 174 variable sites that generated 231 haplotypes were observed. They defined two haplogroups that were present in all the 13 study populations. Reference haplotypes from the six globally defined goat mtDNA haplogroups show the two haplogroups present in Ethiopia to be A and G, the former being the most predominant. Although both haplogroups are characterized by an increase in effective population sizes (N_e) predating domestication, they also have experienced a decline in N_e at different time periods, suggesting different demographic histories. We observed seven haplotypes, six were directly linked to the central haplotypes of the two haplogroups and one was central to haplogroup G. The seven haplotypes were common between Ethiopia, Kenya, Egypt, and Saudi Arabia populations, suggesting common maternal history and the introduction of goats into East Africa via Egypt and the Arabian Peninsula, respectively. While providing new mtDNA data from a historically important region, our results suggest extensive intermixing of goats mediated by human socio‐cultural and economic interactions. These have led to the coexistence of the two haplogroups in different geographic regions in Ethiopia resulting in a large caprine genetic diversity that can be exploited for genetic improvement.     

Ethiopians and Khoisan Share the Deepest Clades of the Human Y-Chromosome Phylogeny

The genetic structure of 126 Ethiopian and 139 Senegalese Y chromosomes was investigated by a hierarchical analysis of 30 diagnostic biallelic markers selected from the worldwide Y-chromosome genealogy. The present study reveals that (1) only the Ethiopians share with the Khoisan the deepest human Y-chromosome clades (the African-specific Groups I and II) but with a repertoire of very different haplotypes; (2) most of the Ethiopians and virtually all the Senegalese belong to Group III, whose precursor is believed to be involved in the first migration out of Africa; and (3) the Ethiopian Y chromosomes that fall into Groups VI, VIII, and IX may be explained by back migrations from Asia. The first observation confirms the ancestral affinity between the Ethiopians and the Khoisan, which has previously been suggested by both archaeological and genetic findings.     

Population genetics of Glossina morsitans submorsitans (Diptera: Glossinidae)

Breeding structure of Glossina morsitans submorsitans Newstead was evaluated by using genetic markers in mitochondrial DNA where diversity was scored at two loci in five natural populations from The Gambia and two populations in Ethiopia (form ugandensis Vanderplank), countries separated by c. 5450 km. Twenty six haplotype combinations were found, of which 17 were shared among two or more populations. Nine haplotypes were found in The Gambia and 23 haplotypes in Ethiopia. There were 12 unique haplotypes. Only six haplotypes were shared between the two countries. Populations in The Gambia (he = 0.26 +/- 0.04) showed less than a third of the diversity of populations in Ethiopia (he = 0.84 +/- 0.03). This suggests recovery from an earlier reduction in population. In a nested analysis of molecular variance of haplotype frequencies, 65% of the variance was due to differences within populations, 34% to differences between populations grouped by country, and only 1% was due to differences among populations within countries. In terms of gene flow, the fixation index FST = 0.35, which leads to an estimate by Wright's island model of less than one reproducing migrant per generation exchanged between the eastern and western submorsitans populations. Nei's genetic similarity measure showed a deep division between Gambian and Ethiopian populations.     

Chloroplast DNA microsatellite analysis supports a polyphyletic origin for barley

Five barley chloroplast DNA microsatellites (cpSSRs) were used to study genetic relationships among a set of 186 barley accessions—34 Hordeum vulgare ssp. spontaneum (HS accessions) from Morocco, Ethiopia, Cyprus, Crete, Libya, Iraq, Iran, Turkey, Afghanistan and Israel, 122 H. vulgare ssp. vulgare landraces (HV landraces) from Spain, Bolivia (old Spanish introductions), Morocco, Libya and Ethiopia and 20 modern European spring barleys (HV cultivars). All loci were polymorphic in the material studied, with the number of alleles per locus ranging from two to three. Fifteen multi-locus haplotypes were observed, 11 in HS accessions and seven in HV landraces and cultivars. Of the seven haplotypes found in the HV lines, three were shared with the HS accessions, and four were unique. Cluster analysis revealed two main groups, one consisting of HS accessions from Ethiopia and the HV landraces from Spain, Bolivia (old Spanish), Morocco and Ethiopia, whereas the other larger group contained all of the other accessions studied. Based on these grouping and the existence of haplotypes found in the HV landraces and cultivars but not in the HS wild barley, a polyphyletic origin is proposed for barley, with further centres of origin in Ethiopia and the Western Mediterranean.     

Ethnic Pluralism as an Organizing Principle of the Ethiopian Federation

In 1991 the Ethiopian government employed ethnic pluralism as an organizing principle, creating multiple ethnic-based territorial units with a right of secession provision. Ethiopians are watching this experiment with considerable apprehension. This paper: (1) provides a concise historical background of ethnic relations in Ethiopia, (2) examines the type of ethnic federal state established in Ethiopia, and (3) points out some problems encountered with ethnicity as an organizing principle and attempts a preliminary assessment of the ethnic-based federal experiment. The sources of data for this paper include public documents, fieldwork, and interviews with 30 knowledgeable Ethiopians in Addis Ababa. I spent several months during 2000–2002 observing political developments in the country as they pertain to ethnic federalism. Within Africa, a nation-state fully acknowledging and based on ethnic pluralism is unique. Thus far, the Ethiopian federation appears to have undercut the drive for secession by largely removing its rallying cause, manifest ethnic oppression. Nonetheless, the fact that the ruling group comes predominantly from a small ethnic group has raised serious protest from other ethnic groups, larger and smaller. Its use of democratic centralism has also undermined effective decentralization and democratization. Ethnic pluralism as an organizing principle underpinning the federal government in Ethiopia is a fragile and perilous experiment.     

Gm and Inv allotypes of some Sidamo Ethiopians

One hundred and forty serum samples from Ethiopians from three tribes of the Sidamo group were tested for their Gm and Inv phenotypes. The Gm antigens 1,2,3,5,6,13,14,21 were determined on all samples, and Gm(15) and Gm(16) were determined on selected samples. All samples were tested for Inv(1), and those positive for Inv(1) were tested for Inv(3). The samples fall into 18 Gm phenotypes and require seven haplotypes to explain them. The data indicate that these Ethiopians have Negroid, Caucasoid, and Bushmanoid ancestry, with the latter constituting a relatively small proportion of the ancestry and the former two contributing about equally to the remainder. The data are consistent with the conclusions of cultural anthropologists.     

Variation in mitochondrial DNA and maternal genetic ancestry of Ethiopian cattle populations

This study describes complete control region sequences of mitochondrial DNA (mtDNA) from 117 Ethiopian cattle from 10 representative populations, in conjunction with the available cattle sequences in GenBank. In total, 79 polymorphic sites were detected, and these defined 81 different haplotypes. The haplotype and nucleotide diversity of Ethiopian cattle did not vary among the populations studied. All mtDNA sequences from Ethiopian cattle converged into one main maternal lineage (T1) that corresponds to African Bos taurus cattle. According to the results of this study, no zebu mtDNA haplotypes have been found in Ethiopia, where the most extensive hybridization took place on the African continent.     

Combined use of biallelic and microsatellite Y-chromosome polymorphisms to infer affinities among African populations.

To define Y-chromosome haplotypes, we studied seven biallelic polymorphic sites. We combined data with those from four dinucleotide-repeat polymorphisms, to establish Y-chromosome compound superhaplotypes. Eight biallelic haplotypes that matched the dendrogram proposed by other investigators were identified in 762 Y chromosomes from 25 African populations. For each biallelic site, coalescence time of lineages carrying the derived allele was estimated and compared with previous estimates. The "ancestral" haplotype (haplotype 1A) was observed among Ethiopians, "Khoisan" (!Kung and Khwe), and populations from northern Cameroon. Microsatellite distributions within this haplotype showed that the Khoisan haplotypes 1A are widely divergent from those of the other two groups. Populations from northern Africa and northern Cameroon share a haplotype (i.e., 1C), which is not observed in other African populations but represents a major Eurasian cluster. Haplotypes 1C of northern Cameroon are clearly distinct from those of Europe, whereas haplotypes 1C of northern African are well intermingled with those of the other two groups. Apportionment of diversity for the Y-chromosomal biallelic haplotypes was calculated after populations were clustered into different configurations. Despite some correspondence between language affiliation and genetic similarity, geographic proximity seems to be a better predictor of genetic affinity.     

Micro- and macrogeographical genetic structure of colonies of naked mole-rats Heterocephalus glaber

Patterns of genetic structure in eusocial naked mole-rat populations were quantified within and among geographically distant populations using multilocus DNA fingerprinting and mitochondrial DNA (mtDNA) sequence analysis. Individuals within colonies were genetically almost monomorphic, sharing the same mtDNA control region haplotype and having coefficients of band sharing estimated from DNA fingerprints ranging from 0.93 to 0.99. Family analysis of a hybrid captive colony of naked mole-rats with increased levels of genetic variability using multilocus DNA fingerprinting gave results consistent with Mendelian inheritance, and has revealed for the first time that multiple paternity can occur. In a survey of wild colonies from Ethiopia, Somalia and locations in northern and southern Kenya, we have examined mtDNA control region sequence variation in 42 individuals from 15 colonies, and together with multilocus DNA fingerprinting and mtDNA cytochrome- b sequence analysis in selected individuals have shown that these populations show considerable genetic divergence. Most of the variance in sequence divergence was found to be between geographical locations (Φ_ct= 0.68) and there was a significant correlation between sequence divergence and geographical separation of haplotypes. Six colonies from Mtito Andei in southern Kenya shared the same control region haplotype, suggesting a recent common maternal ancestor. In contrast, out of four colonies at Lerata in north Kenya, three haplotypes were identified, and phylogenetic analysis suggests that this area may be a zone where two distinct lineages are in close proximity. Genetic distances were maximal between Ethiopian and southern Kenyan populations at 5.8% for cytochrome- b , and are approaching interspecific values seen between other Bathyergids.     

Molecular identification of unilocular hydatid cysts from domestic ungulates in Ethiopia: implications for human infections

To identify the etiologic agents of cystic echinococcosis in Ethiopia, unilocular hydatid cysts were collected from 11 sheep, 16 cattle and 16 camels slaughtered in abattoirs of Aweday, Jijiga, Haramaya and Addis Ababa during June 2010 to February 2011. A PCR-based DNA sequencing of the mitochondrial cytochrome oxidase c subunit 1 gene (cox1) was conducted for 40 cysts. The majority of cysts (87.5%) were identified as Echinococcus granulosus sensu stricto and the rest as Echinococcus canadensis. The fertile cysts of E. granulosus s.s. were found only from sheep, although it occurred in all the host species. The predominance of E. granulosus s.s. has important implications for public health since this species is the most typical causative agent of human cystic echinococcosis worldwide. The major cox1 haplotype of E. granulosus s.s. detected in Ethiopia was the same as that has been reported to be most common in Peru and China. However, a few cox1 haplotypes unique to Ethiopia were found in both of the two Echinococcus species. The present regional data would serve as baseline information in determining the local transmission patterns and in designing appropriate control strategies.     

Tracing the Route of Modern Humans out of Africa by Using 225 Human Genome Sequences from Ethiopians and Egyptians

The predominantly African origin of all modern human populations is well established, but the route taken out of Africa is still unclear. Two alternative routes, via Egypt and Sinai or across the Bab el Mandeb strait into Arabia, have traditionally been proposed as feasible gateways in light of geographic, paleoclimatic, archaeological, and genetic evidence. Distinguishing among these alternatives has been difficult. We generated 225 whole-genome sequences (225 at 8× depth, of which 8 were increased to 30×; Illumina HiSeq 2000) from six modern Northeast African populations (100 Egyptians and five Ethiopian populations each represented by 25 individuals). West Eurasian components were masked out, and the remaining African haplotypes were compared with a panel of sub-Saharan African and non-African genomes. We showed that masked Northeast African haplotypes overall were more similar to non-African haplotypes and more frequently present outside Africa than were any sets of haplotypes derived from a West African population. Furthermore, the masked Egyptian haplotypes showed these properties more markedly than the masked Ethiopian haplotypes, pointing to Egypt as the more likely gateway in the exodus to the rest of the world. Using five Ethiopian and three Egyptian high-coverage masked genomes and the multiple sequentially Markovian coalescent (MSMC) approach, we estimated the genetic split times of Egyptians and Ethiopians from non-African populations at 55,000 and 65,000 years ago, respectively, whereas that of West Africans was estimated to be 75,000 years ago. Both the haplotype and MSMC analyses thus suggest a predominant northern route out of Africa via Egypt.     

Pleistocene colonization of afro-alpine 'sky islands' by the arctic-alpine Arabis alpina

The afro-alpine region comprises the high mountains of Ethiopia and tropical East Africa, which represent biological 'sky islands' with high level of endemism. However, some primarily arctic-alpine plants also occur in the afro-alpine mountains. It has been suggested that these plants are Tertiary relicts, but a recent worldwide study of Arabis alpina suggests that this species colonized the region twice during the Pleistocene. Here we investigate the detailed colonization history of A. alpina in the afro-alpine region based on chloroplast DNA sequences from 11 mountain systems. The results confirm the twice-into-Africa scenario. The Asian lineage is confined to the mountains closest to the Arabian Peninsula, on opposite sides of the Rift Valley (Simen Mts and Gara Muleta in Ethiopia), suggesting long-distance dispersal of this lineage. The African lineage is divided into two phylogeographic groups with distinct geographic distribution. The observed pattern is consistent with isolation of the African lineage in at least two interglacial refugia, located on separated highlands, followed by range expansion in cooler period(s), when the afro-alpine habitat extended further down the mountains. Several long-distance dispersal events, also across the Rift Valley, are suggested by single haplotypes observed outside the area occupied by the phylogeographic groups they belonged to.     

Ethiopian mitochondrial DNA heritage: tracking gene flow across and around the gate of tears

Approximately 10 miles separate the Horn of Africa from the Arabian Peninsula at Bab-el-Mandeb (the Gate of Tears). Both historic and archaeological evidence indicate tight cultural connections, over millennia, between these two regions. High-resolution phylogenetic analysis of 270 Ethiopian and 115 Yemeni mitochondrial DNAs was performed in a worldwide context, to explore gene flow across the Red and Arabian Seas. Nine distinct subclades, including three newly defined ones, were found to characterize entirely the variation of Ethiopian and Yemeni L3 lineages. Both Ethiopians and Yemenis contain an almost-equal proportion of Eurasian-specific M and N and African-specific lineages and therefore cluster together in a multidimensional scaling plot between Near Eastern and sub-Saharan African populations. Phylogeographic identification of potential founder haplotypes revealed that approximately one-half of haplogroup L0–L5 lineages in Yemenis have close or matching counterparts in southeastern Africans, compared with a minor share in Ethiopians. Newly defined clade L6, the most frequent haplogroup in Yemenis, showed no close matches among 3,000 African samples. These results highlight the complexity of Ethiopian and Yemeni genetic heritage and are consistent with the introduction of maternal lineages into the South Arabian gene pool from different source populations of East Africa. A high proportion of Ethiopian lineages, significantly more abundant in the northeast of that country, trace their western Eurasian origin in haplogroup N through assorted gene flow at different times and involving different source populations.     

Mitochondrial DNA and Y‐chromosome diversity in East Adriatic sheep

Variation in mitochondrial DNA (mtDNA) and Y‐chromosome haplotypes was analysed in nine domestic sheep breeds (159 rams) and 21 mouflon ( Ovis musimon) sampled in the East Adriatic. Mitochondrial DNA analyses revealed a high frequency of type B haplotypes, predominantly in European breeds, and a very low frequency of type A haplotypes, which are more frequent in some Asian breeds. Mitochondrial haplotype Hmt‐3 was the most frequent (26.4%), and 37.1%, 20.8% and 7.6% of rams had haplotypes one, two and three mutations remote from Hmt‐3 respectively. In contrast, Y‐chromosome analyses revealed extraordinary paternal allelic richness: HY‐6, 89.3%; HY‐8, 5.0%; HY‐18, 3.1%; HY‐7, 1.3%; and HY‐5, 1.3%. In fact, the number of haplotypes observed is comparable to the number found in Turkish breeds and greater than the number found in European breeds so far. Haplotype HY‐18 (A‐oY1/135‐SRYM18), identified here for the first time, provides a link between the haplotype HY‐12 (A‐oY1/139‐SRYM18) found in a few rams in Turkey and haplotype HY‐9 (A‐oY1/131‐SRYM18) found in one ram in Ethiopia. All mouflons had type B mtDNA haplotypes, including the private haplotype (Hmt‐55), and all were paternally monomorphic for haplotype HY‐6. Our data support a quite homogeneous maternal origin of East Adriatic sheep, which is a characteristic of European breeds. At the same time, the high number of haplotypes found was surprising and intriguing, and it begs for further analysis. Simultaneous analysis of mtDNA and Y‐chromosome information allowed us to detect a large discrepancy between maternal and paternal lineages in some populations. This is most likely the result of breeder efforts to ‘upgrade’ local populations using rams with different paternal origins.     

Phantom Wars and Cyberwars: Abyssinian Fundamentalism and Catastrophe in Eritrea

The 1998–2000 war between Ethiopia and Eritreapitted former allies against each other, endingnew opportunities for development anddemocracy. We examine discursive constructionof the war, including Western media coverageand propaganda from both governments, but focuson discourse of exiled Ethiopians committed topolitical struggles in their former homeland,using the concept of long-distance nationalism.To investigate one form of long-distancenationalism, resurgent Abyssinianfundamentalism, we turn to sites of itsdiscursive production, including the magazine,Ethiopian Review, and the Internet. Theorists of transnationalism have consideredthe Internet as a site for new forms ofcommunication and community and argue thatInternet discourse among diaspora professionalsand academics embodies a creolizeddiscourse produced by a distinctive type, thecybernaut. In the case of the Abyssiniancybernauts, however, we detect not a newcreolized discourse but a ghostly repetitionof old views.     

Phylogeography of the heathers Erica arborea and E. trimera in the afro-alpine ‘sky islands’ inferred from AFLPs and plastid DNA sequences

The ericaceous vegetation zone of the unique and highly fragmented afro-alpine environment in the eastern African high mountains is typically dominated by the heather Erica arborea, often in combination with its close relative E. trimera. Both species are shrubs or small trees with tiny seeds, potentially capable of dispersal by wind over long distances. While E. arborea is widely distributed in Africa, the Middle East and Europe, E. trimera is endemic to the afro-alpine region where it is restricted to higher altitudes than E. arborea. We used Amplified Fragment Length Polymorphisms (AFLPs) and variation in non-coding plastid DNA sequences to test whether these two morphologically and ecologically very similar species display similar phylogeographic patterns in the afro-alpine region. We predict that the more high-altitudinal E. trimera shows more distinct genetic structuring than E. arborea, because dispersal of the latter may have been facilitated by formation of interglacial forest bridges between mountains. Based on extensive field sampling in most of the high mountains of Ethiopia and East Africa, we show that the two species are clearly distinct at AFLP and plastid DNA loci. Both showed low levels of overall AFLP diversity, suggesting bottlenecking in small refugial populations during unfavourable climatic periods. However, their genetic structuring and inferred phylogeographic histories were conspicuously different. The more high-altitudinal E. trimera consisted of three to four distinct AFLP groups, which also had different plastid DNA haplotypes and different geographic distributions, suggesting long-term restriction to several refugia (at least one in Ethiopia and two in East Africa). In contrast, E. arborea showed little geographic structuring at AFLP loci and only a single, widespread plastid DNA haplotype, which may suggest recent colonization of the entire study area from a single source population, likely via a combination of gradual expansion via forest bridges and long-distance dispersals. The source population of E. arborea may be situated in (or north of) Ethiopia, which harbours most genetic diversity.     

Gm and Inv studies on baboons, Papio cynocephalus: analysis of serum samples from Kenya, Ethiopia and South Africa.

Serum samples from 526 baboons (Papio cynocephalus) from 10 troops from the Laikipia district of northern Kenya, from 60 baboons from two troops from the Awash National Park, central Ethiopia, and from 127 baboons from South Africa were tested for Gm and Inv allotypes. Four of the 10 troops from Kenya formed a western cluster and six formed an eastern cluster. The clusters were separated by approximately 10 miles. The samples were tested for Gm (1, 2, 3, 5, 6, 11, 13, 14, 15, 16, 17, 21, 24) and for Inv (1, 2, 3). All samples were negative for Gm (2, 6, 14, 16, 24). All from Kenya and Ethiopia were negative for Inv (2), and all were positive for Gm (11, 17) and for Inv (3). The south African samples differed from the others in that 10 were negative for Gm (11) and four were positive for Inv (2). Taking all animals into account, polymorphism was present for Gm (1, 3, 5, 11, 13, 15, 21) and for Inv (1, 2). No two Kenya troops had the same array of phenotypes or of haplotypes, but the four western troops were more similar to each other than to the six eastern troops. Three haplotypes were present in the eastern troops that were not present in the western troops and five were present in the western troops that were not present in the eastern troops. Five haplotypes appeared in at least some troops of each cluster of troops. The samples from each of the two troops from Ethiopia show the same three phenotypes but with significantly different frequencies. It is suggested that the variation in haplotype frequencies observed among the 10 troops from Kenya is the result of a founder effect deriving largely from fission of a large troop into two smaller troops. The data show that speculations about the evolutionary origin of the allotypes are premature. For most species, too few animals have been tested and except for those in this study their origins are not known. Finally, the samples have been from too restricted an area.