Domestication ofEleusine coracana

Eleusine coracana (L.) Gaertn. subsp.coracana (Finger millet or eleusine) is a cereal that is widely cultivated in Africa and India. Archaeological records of finger millet are few and unsatisfactory. But, distribution, linguistic and historical evidence seem to suggest an African rather than Indian origin of the crop. Data from morphology, supplemented with cytogenetical observations and distribution revealed thatE. coracana subsp.africana is wild finger millet. This subspecies is widely distributed along the highlands of East Africa. Consequently, it is concluded that finger millet originated in the East African Highlands and was subsequently introduced into India.     

SYSTEMATICS AND EVOLUTION OF ELEUSINE CORACANA (GRAMINEAE)

Finger millet (Eleusine coracana (L.) Gaertn. subsp. coracana) is cultivated in eastern and southern Africa and in southern Asia. The closest wild relative of finger millet is E. coracana subsp. africana (Kennedy-O'Byrne) Hilu & de Wet. Wild finger millet (subsp. africana) is native to Africa but was introduced as a weed to the warmer parts of Asia and America. Derivatives of hybrids between subsp. coracana and subsp. africana are companion weeds of the crop in Africa. Cultivated finger millets are divided into five races on the basis of inflorescence morphology. Race coracana is widely distributed across the range of finger millet cultivation. It is present in the archaeological record of early African agriculture that may date back 5,000 years. Racial evolution took place in Africa. Races vulgaris, elongata, plana, and compacta evolved from race coracana, and were introduced into India some 3,000 years ago. Little independent racial evolution took place in India.     

ARCHAEOBOTANICAL STUDIES OF ELEUSINE CORACANA SSP. CORACANA (FINGER MILLET)

Two controversial archaeological specimens of finger millet, Eleusine coracana ssp. coracana (L.) Gaertn., were examined by light and scanning electron microscopy. Identification of the material was based on morphological and anatomical comparison with laboratory carbonized grains of modern Eleusine. Seed morphology of the Indian archaeological collection did not conform with those of domesticated or wild species of Eleusine. The other archaeological collection examined is from Ethiopia. It is estimated to date back to the third millennium B.C. These well preserved inflorescence fragments were positively identified as finger millet. If the suggested date is correct, they represent the oldest archaeological record for the crop, as well as the oldest agricultural record for Africa south of the Sahara. These findings support biosystematic evidence for an East African origin of finger millet which leaves India as a secondary center of diversity.     

Population Structure and Diversity in Finger Millet (Eleusine coracana) Germplasm

A genotypic analysis of 79 finger millet accessions (E. coracana subsp. coracana) from 11 African and five Asian countries, plus 14 wild E. coracana subsp. africana lines collected in Uganda and Kenya was conducted with 45 SSR markers distributed across the finger millet genome. Phylogenetic and population structure analyses showed that the E. coracana germplasm formed three largely distinct subpopulations, representing subsp. africana, subsp. coracana originating from Africa and subsp. coracana originating from Asia. A few lines showed admixture between the African and Asian cultivated germplasm pools and were the result of either targeted or accidental intercrossing. Evidence of gene flow was also seen between the African wild and cultivated subpopulations, indicating that hybridizations among subspecies occur naturally where both species are sympatric. The genotyping, combined with phylogenetic and population structure analyses proved to be very powerful in predicting the origin of breeding materials. The genotypic study was complemented by a phenotypic evaluation. The wild and cultivated accessions differed by a range of domestication-related characters, such as tiller number, plant height, peduncle length, seed color and grain yield. Significant differences in plant architecture and yield were also identified between the Asian and African subpopulations. The observed population structure within cultivated finger millet is consistent with the theory that, after the introduction of finger millet from Africa into India via the trade routes some 3000 years ago, the two germplasm pools remained largely isolated until recent times. The significantly lower diversity present within the Asian subpopulation also suggests that it arose from a relatively small number of founder plants.     

Flavonoid patterns and systematics in Eleusine

A survey of leaf flavonoids was conducted on Eleusine coracana ssp. coracana and ssp. africana, E. indica, E. multiflora, E. tristachya, E. floccifolia, and E. compressa. Twenty phenolic compounds were detected. Those identified were: orientin, isoorientin, vitexin, isovitexin, saponarin, violanthin, lucenin-1, and tricin. The study revealed a general generic flavonoid pattern except for E. compressa, which occupies an isolated position in Eleusine. Flavonoids of the perennial E. floccifolia and the annuals E. multiflora and E. tristachya are markedly different from those of cultivated E. coracana, suggesting that these species are only distantly related to the crop. The morphologically well defined E. coracana—africana—indica group also forms a unit in respect of flavonoids. Subspecies africana exhibits a higher flavonoid similarity to ssp. coracana (finger millet) than does E. indica. The weedy race of ssp. africana usually combines flavonoids of both the wild and domesticated subspecies. The flavonoid pattern of the dedza race of ssp.africana is identical to that of finger millet, suggesting either a direct origin of the crop from this race, or extensive introgression from the crop into ssp. africana. A lack of qualitative differences in flavonoids between cultivated races of finger millet is indicative of the genetic stability of these compounds. The flavonoid data confirms the domestication of finger millet from ssp. africana.     

A historical overview of the appearance and spread of Musa pests and pathogens on the African continent: highlighting the importance of clean Musa planting materials and quarantine measures

The genus Musa is not native to Africa. It evolved in tropical Asia, from southwest India eastward to the island of New Guinea. There is a growing circumstantial evidence which suggests that the East African Highland banana and the tropical lowland plantain were cultivated on the African continent since before 1 AD. It is also probable that ABB cooking and AB and AAB dessert cultivars were brought to the continent from India by Arabian traders from 600 AD, and that these were disseminated throughout East Africa. During the colonial era, the main centres of distribution for banana cultivars were botanical gardens, such as Zomba in Malawi, Entebbe in Uganda and Amani in Tanzania. It appears that the very early introductions of Highland banana and plantain arrived in Africa as a relatively clean material without the conspicuous pests and diseases that affect them in Asia. In contrast, several devastating problems now impact the crop in Africa, including nematodes, the borer weevil and diseases, most notably banana bunchy top, banana streak, Sigatoka leaf spots, Xanthomonas wilt and Fusarium wilt. We (a) provide chronological overviews of the first reports/observations of different Musa pests and pathogens/diseases in Africa, (b) highlight specific examples of when a pest or pathogen/disease was introduced via planting materials and (c) give recent examples of how the pests and pathogens spread to new regions via planting materials. In total, these production constraints threaten banana and plantain production throughout the continent and impact those who can ill afford lost production, the small‐holder producer. Our intent in this review is to highlight the significance of these problems and the great importance that infested planting materials have played in their development.     

Pathogenic Variability in Colletotrichum lindemuthianum and Evaluation of Resistance in Phaseolus vulgaris in the North-Western Himalayan Region of India

Eightyfive isolates of Colletotrichum lindemuthianum (bean anthracnose) collected from different kidney bean growing areas of a hilly state (Himachal Pradesh) of India, were characterized on the basis of their reaction types on International and CIAT differentials. On international differentials, 12 races viz., Alpha-Brazil, Beta, Gamma and Ind I to Ind IX were characterized. The races designated as Ind I to Ind IX were different from those identified in Europe and USA, thus forming a new race group from the Indian subcontinent. On the CIAT differential set the 85 isolates have been grouped into 19 races. Of these, only races 65 and 73 resembled the North American races. Exotic accessions AB 136 and G 2333 were resistant to all the Indian races. However, race specific resistance has been found in a number of indigenous and exotic genotypes of Phaseolus vulgaris.     

Race Profiling and Molecular Diversity Analysis of Fusarium oxysporum f.sp. ciceris Causing Wilt in Chickpea

Seventy isolates of Fusarium oxysporum f.sp. ciceris (Foc) causing chickpea wilt representing 13 states and four crop cultivation zones of India were analysed for their virulence and genetic diversity. The isolates of the pathogen showed high variability in causing wilt incidence on a new set of differential cultivars of chickpea, namely C104, JG74, CPS1, BG212, WR315, KWR108, GPF2, DCP92‐3, Chaffa and JG62. New differential cultivars for each race were identified, and based on differential responses, the isolates were characterized into eight races of the pathogen. The same set of isolates was used for molecular characterization with four different molecular markers, namely random amplified polymorphic DNA, universal rice primers, simple sequence repeats and intersimple sequence repeats. All the four sets of markers gave 100% polymorphism. Unweighted paired group method with arithmetic average analysis grouped the isolates into eight categories at genetic similarities ranging from 37 to 40%. The molecular groups partially corresponded to the states of origin/chickpea‐growing region of the isolates as well as races of the pathogen characterized in this study. The majority of southern, northern and central Indian populations representing specific races of the pathogen were grouped separately into distinct clusters along with some other isolates, indicating the existence of variability in population predominated by a single race of the pathogen. The present race profiling for the Indian population of the pathogen and its distribution pattern is entirely new. The knowledge generated in this study could be utilized in resistance breeding programme. The existence of more than one race, predominated by a single one, in a chickpea cultivation zone as supported by the present molecular findings is also a new information.     

The linyphiid fauna of eastern Africa (Araneae: Linyphiidae)—distribution patterns, diversity and endemism

Distributions, endemism and diversity among East African linyphiids are analysed and discussed in relation to other forest organisms and the environmental history of eastern African. A total of 231 species are reported from eastern Africa, of which 14 are confined to the Afroalpine region and 114 species to moist forests. Only 12 of the latter are widely distributed. The rest are only known from one or two localities. Information on habitats and distributions of all species is tabulated. Few species are shared between East African mountains and there are no detectable gradients of species diversity between mountains. There is, however, a gradient of decreasing species diversity from high latitudes to the Equator. Vicariance patterns are demonstrated for Elgonia, Ophrynia and Callitrichia in the Eastern Arc Mountains, Tanzania—areas that also hold the highest degree of endemism (> 80% on individual mountains) among linyphiids. The many endemic species on nearby mountains suggest that intermontane dispersal (ballooning) is rare or non-existent. There is no evidence for a distinction between highland and lowland linyphiid faunas, but altitudinal segregation of single species is demonstrated. The question of the reality of highland and lowland faunas cannot be solved by studying the altitudinal distribution of single species. Phylogenetic relationships must be taken into consideration to determine where sister-groups/species are located (lowland or highland).     

Projecting the effects of climate change on the distribution of maize races and their wild relatives in Mexico

Climate change is expected to be a significant threat to biodiversity, including crop diversity at centers of origin and diversification. As a way to avoid food scarcity in the future, it is important to have a better understanding of the possible impacts of climate change on crops. We evaluated these impacts on maize, one of the most important crops worldwide, and its wild relatives Tripsacum and Teocintes. Maize is the staple crop in Mexico and Mesoamerica, and there are currently about 59 described races in Mexico, which is considered its center of origin . In this study, we modeled the distribution of maize races and its wild relatives in Mexico for the present and for two time periods in the future (2030 and 2050), to identify the potentially most vulnerable taxa and geographic regions in the face of climate change. Bioclimatic distribution of crops has seldom been modeled, probably because social and cultural factors play an important role on crop suitability. Nonetheless, rainfall and temperature still represent a major influence on crop distribution pattern, particularly in rainfed crop systems under traditional agrotechnology. Such is the case of Mexican maize races and consequently, climate change impacts can be expected. Our findings generally show significant reductions of potential distribution areas by 2030 and 2050 in most cases. However, future projections of each race show contrasting responses to climatic scenarios. Several evaluated races show new potential distribution areas in the future, suggesting that proper management may favor diversity conservation. Modeled distributions of Tripsacum species and Teocintes indicate more severe impacts compared with maize races. Our projections lead to in situ and ex situ conservation recommended actions to guarantee the preservation of the genetic diversity of Mexican maize.     

Pathogen race determines the type of resistance response in the stripe rust –Triticum dicoccoides pathosystem

Wild relatives of crop plants may serve as a promising source for screening for new disease resistance genes that can be utilized in breeding programs. Triticum dicoccoides, the wild progenitor of most cultivated wheats, was shown to harbor many resistance genes against the major diseases attacking cultivated wheat. Stripe rust is a devastating fungal disease that attacks wheat in many regions of the world. New races of Puccinia striiformis Westend. f. sp. tritici, the causative agent of stripe rust, have overcome most of the known Yr resistance genes in wheat. Therefore, there is a need to search for new resistance genes in the T. dicoccoides gene pool. A set of 120 T. dicoccoides accessions, collected from 13 populations representing different habitats in Israel and vicinity, was tested for resistance to three prevalent stripe rust races (38E134, 6E16 and 6E0). Of these 120 accessions, 14, 8 and 12% were resistant to races 38E134, 6E16 and 6E0, respectively, while 57, 2 and 4% were moderately resistant to these races, respectively. A unique resistance was found in the population of Mt Hermon where >80% of the accessions showed resistance to all races. Distribution of infection types (ITs) of race 38E134 showed a normal distribution that can fit a quantitative pattern of response, while the distributions of ITs of races 6E16 and 6E0 had excess of extreme values and therefore showing a qualitative pattern of response. anova testing the main factor effects and interaction showed significant effects of population, race and their interaction on IT. Significant positive correlations were obtained between the resistance to races 6E16 and 6E0 and humidity variables of the collections sites, while resistance to race 38E134 was positively correlated with temperature variables. These results show that the pathogen race can determine the type of resistance response, qualitative or quantitative, in the stripe rust—T. dicoccoides pathosystem. The obtained results also reveal that the distribution of resistance to different pathogen races can be affected by different climatic factors.     

Distribution and biogeography of the Paradiaptominae (Copepoda: Calanoida: Diaptomidae)

The Paradiaptominae, a subfamily of the large Diaptomidae family, is a small group of freshwater calanoids of which 24 species in four genera have been described. They are endemic to Africa with the exception of four species, Metadiaptomus asiaticus (Asia and Mongolia), M. gauthieri (Africa and Madagascar), Paradiaptomus greeni (India and Sri Lanka) and Neolovenula alluaudi (Canary Islands, North Africa, Baltic and Mediterranean countries). The North African species Metadiaptomus chevreuxi extends its range into Iraq and Iran. The Paradiaptominae are widely distributed in semi-permanent water bodies in the drier areas of Africa although most individual species have a restricted distribution. Only a few species have the ability to colonise permanent waters and these species are usually located at higher altitudes. The Paradiaptominae have not been recorded from the wet equatorial lowland areas of Africa. It seems likely that they are Gondwanian in origin as evidenced by their distribution (Africa, Madagascar, India and Sri Lanka). Neolovenula alluaudi (Mediterranean) shares morphological characters tenuously linked to the other species in the group and may either have a different origin or have branched off at an early stage in evolution of the group.     

RFLP diversity in cultivated sorghum in relation to racial differentiation

Careful assessment of the comparative diversity for molecular markers and for potentially-useful morpho-agronomic traits is paramount to the analysis of a genome through the mapping of favorable genes. Sorghum (Sorghum bicolor ssp.bicolor) varieties are traditionally classified into five races on the basis of morphological traits, especially panicle and grain traits. Isozyme diversity has provided a new insight into genetic diversity, and showed a marked geographic structure. We performed RFLP analysis on 94 varieties, chosen to represent the main cross combinations (race × geographic origin), using 35 maize probes that detect polymorphism with at least one of the two restriction enzymesHindIII andXbaI. A total of 50 polymorphic probe-enzyme combinations yielded 158 polymorphic bands. The bicolor race appeared highly variable and included many rare markers. Among the other races multivariate analysis of the data differentiated six clusters corresponding, by decreasing magnitude of divergence, to: the margaritiferum types (a sub-race of race guinea); the guinea forms from western Africa; race caudatum; race durra; race kafir; and the guinea forms from southern Africa.The apparent geographic differentiation was related to the contrasting distribution of these races and to a higher similarity between races localized in southern Africa. The data agree with the current hypotheses on sorghum domestication but reveal associations between neutral markers and traits probably highly subjected to human selection. Whether such associations will be observed with other useful traits, and to what extent they are maintained by genetic linkage, is worth exploring.     

Use of genomic and genic SSR markers for assessing genetic diversity and population structure in Indian and African finger millet (Eleusine coracana (L.) Gaertn.) germplasm

Genetic diversity and population structure were analyzed in 67 diverse finger millet accessions of African and Indian origin. A total of 69 alleles were generated with a mean of 4.0 alleles per locus and with an average gene diversity of 0.471. Molecular diversity parameters showed higher values in African accessions. Nineteen rare and nine unique alleles were observed and the identified accessions can be a potential source for further improvement of finger millet. Clustering of south Indian accessions with African lowland types and north/highland Indian accessions with that of African highlands was also observed. Structure analysis revealed the distinctness of Ugandan accessions compared to other African accessions and five major sub-populations useful for parental selection, conservation, and utilization of finger millet.     

SPECIAL PAPER: SYSTEMATICS AND EVOLUTION OF SORGHUM SECT. SORGHUM (GRAMINEAE)

The genus Sorghum Moench is subdivided into sections Chaeotosorghum, Heterosorghum, Parasorghum, Stiposorghum and Sorghum. Section Sorghum includes two rhizomatous species, S. halepense (L.) Pers. (2n = 40) and S. propinquum (Kunth) Hitchcock (2n = 20), as well as the annual S. bicolor (L.) Moench (2n = 20). Sorghum bicolor is divided into subspecies bicolor to include all domesticated grain sorghums, subspecies drummondii (Steud.) de Wet comb. nov. to include stabilized derivatives of hybridization among grain sorghums and their closest wild relatives and subspecies arundinaceum (Desv.) de Wet et Harlan to include the wild progenitors of grain sorghums. Four ecotypes of subspecies arundinaceum are recognized: race aethiopicum of the arid African Sahel. race virgatum of northeastern Africa, race arundinaceum of the African tropical forest, and race verticilliflorum of the African Savanna. The numerous, usually recognized grain sorghums are divided among five basic races, bicolor, caudatum, durra, guinea and kafir, and ten hybrid races that each combine characteristics of at least two of these basic races. Races of grain sorghum are morphologically distinct, and they maintain their unity of type through spacial and ethnological isolation.     

Evolution along the Great Rift Valley: phenotypic and genetic differentiation of East African white‐eyes (Aves,Zosteropidae)

The moist and cool cloud forests of East Africa represent a network of isolated habitats that are separated by dry and warm lowland savannah, offering an opportunity to investigate how strikingly different selective regimes affect species diversification. Here, we used the passerine genus Zosterops (white‐eyes) from this region as our model system. Species of the genus occur in contrasting distribution settings, with geographical mountain isolation driving diversification, and savannah interconnectivity preventing differentiation. We analyze (1) patterns of phenotypic and genetic differentiation in high‐ and lowland species (different distribution settings), (2) investigate the potential effects of natural selection and temporal and spatial isolation (evolutionary drivers), and (3) critically review the taxonomy of this species complex. We found strong phenotypic and genetic differentiation among and within the three focal species, both in the highland species complex and in the lowland taxa. Altitude was a stronger predictor of phenotypic patterns than the current taxonomic classification. We found longitudinal and latitudinal phenotypic gradients for all three species. Furthermore, wing length and body weight were significantly correlated with altitude and habitat type in the highland species Z. poliogaster. Genetic and phenotypic divergence showed contrasting inter‐ and intraspecific structures. We suggest that the evolution of phenotypic characters is mainly driven by natural selection due to differences in the two macro‐habitats, cloud forest and savannah. In contrast, patterns of neutral genetic variation appear to be rather driven by geographical isolation of the respective mountain massifs. Populations of the Z. poliogaster complex, as well as Z. senegalensis and Z. abyssinicus, are not monophyletic based on microsatellite data and have higher levels of intraspecific differentiation compared to the currently accepted species.     

Genome organization and polyploid evolution in the genus Eleusine (Poaceae)

 Eleusine (Poaceae) includes six diploid and three polyploid species and has three basic chromosome numbers, x=8, 9 and 10. The species are annual as well as perennial and all are wild except E. coracana, which is cultivated for grain and fodder in Africa and the Indian subcontinent. Eleusine coracana and E. africana have the same genome and chromosome number (2n=36). Eleusine indica and E. floccifolia are identified as two genome donors to these polyploid species. Eleusine kigeziensis is the third polyploid species of the genus with 2n=38. Its genome may have come from E. jaegeri and from one of the species with x=9, most probably from E. indica. Eleusine indica, E. tristachya, E. floccifolia and E. intermedia with x=9 and two polyploid species, E. coracana and E. africana, are closely related and there is free genetic flow between them. Eleusine multiflora with x=8 is significantly different in morphology and at genomic level from other species. Eleusine jaegeri with x=10 is morphologically similar to E. indica, however, more information is needed to ascertain its position in the genus. Eleusine coracana, which is commonly called finger millet, is a potential and nutritious crop for the increasing population of the world, particularly in arid and semi-arid regions. It can also serve as a gene pool for various important characters and disease resistant genes. Received February 11, 2002; accepted May 27, 2002 Published online: October 14, 2002 Addresses of the authors: Madho Singh Bisht and Yasuhiko Mukai (e-mail: ymukai@cc.osaka-kyoiku.ac.jp), Laboratory of Plant Molecular Genetics, Division of Natural Science, Osaka Kyoiku University, 4-698-1 Asahigaoka, Kashiwara, Osaka 582-8582, Japan.     

Out of Africa? A dated molecular phylogeny of the cicada tribe Platypleurini Schmidt (Hemiptera: Cicadidae), with a focus on African genera and the genus Platypleura Amyot & Audinet‐Serville

The Platypleurini is a large group of charismatic cicadas distributed from Cape Agulhas in South Africa, through tropical Africa, Madagascar, India and eastern Asia to Japan, with generic diversity concentrated in equatorial and southern Africa. This distribution suggests the possibility of a Gondwanan origin and dispersal to eastern Asia from Africa or India. We used a four‐gene (three mitochondrial) molecular dataset, fossil calibrations and molecular clock information to explore the phylogenetic relationships of the platypleurine cicadas and the timing and geography of their diversification. The earliest splits in the tribe were found to separate forest genera in Madagascar and equatorial Africa from the main radiation, and all of the Asian/Indian species sampled formed a younger clade nested well within the African taxa. The tribe appears to have diversified during the Cenozoic, beginning c. 50–32 Ma, with most extant African lineages originating in the Miocene or later, well after the breakup of the Gondwanan landmass. Biogeographical analysis suggests an African origin for the tribe and a single dispersal event founding the Asian platypleurines, although additional taxon sampling and genetic data will be needed to confirm this pattern because key nodes in the tree are still weakly supported. Two Platypleurini genera from Madagascar (Pycna Amyot & Audinet‐Serville, Yanga Distant) are found to have originated by late Miocene dispersal of a single lineage from Africa. The genus Platypleura is recovered as polyphyletic, with Platypleura signifera Walker from South Africa and many Asian/Indian species apparently requiring assignment to different genera, and a new Platypleura concept is proposed with the synonymization of Azanicada Villet syn.n. The genera Orapa Distant and Hamza Distant, currently listed within separate tribes but suspected of platypleurine affinity, are nested deeply within the Platypleurini radiation. The tribe Orapini syn.n . is here synonymized while the tribe Hamzini is pending a decision of the ICZN to preserve nomenclatorial stability.     

Population structure and genetic diversity of New World maize races assessed by DNA microsatellites

Because of the economic importance of maize and its scientific importance as a model system for studies of domestication, its evolutionary history is of general interest. We analyzed the population genetic structure of maize races by genotyping 964 individual plants, representing almost the entire set of ～350 races native to the Americas, with 96 microsatellites. Using Bayesian clustering, we detected four main clusters consisting of highland Mexican, northern United States (US), tropical lowland, and Andean races. Phylogenetic analysis indicated that the southwestern US was an intermediary stepping stone between Mexico and the northern US. Furthermore, southeastern US races appear to be of mixed northern flint and tropical lowland ancestry, while lowland middle South American races are of mixed Andean and tropical lowland ancestry. Several cases of post-Columbian movement of races were detected, most notably from the US to South America. Of the four main clusters, the highest genetic diversity occurs in highland Mexican races, while diversity is lowest in the Andes and northern US. Isolation by distance appears to be the main factor underlying the historical diversification of maize. We identify highland Mexico and the Andes as potential sources of genetic diversity underrepresented among elite lines used in maize breeding programs.