Afrotropical Culicoides : a redescription of C. (Avaritia) imicola Kieffer, 1913 (Diptera : Ceratopogonidae) with description of the closely allied C. (A.) bolitinos sp. nov. reared from the dung of the African buffalo, blue wildebeest and cattle in South Africa

Culicoides (Avaritia) imicola, Kieffer is redescribed and its current worldwide distribution reviewed. It is compared with the closely allied C. (A.) bolitinos sp. nov. Descriptions of both sexes of C. imicola and C. bolitinos sp. nov. are based entirely on series of reared adults. T-tests were performed on antennal and palpal data to differentiate more clearly these 2 species. The results are tabulated. Short notes are given on the larval habitat of C. bolitinos sp. nov. in South Africa, namely the dung of the African buffalo, Syncerus caffer, the blue wildebeest, Connochaetes taurinus, and cattle, Bos races. The existence of C. bolitinos in Nigeria, Kenya, Malawi, Zimbabwe, Lesotho and South Africa is established while C. imicola is newly recorded from Malawi, Botswana, Namibia and Swaziland. Finally seven references in the literature are shown to refer either to C. bolitinos sp. nov. or to closely allied as yet undescribed species.The articles have been scanned in colour with a HP Scanjet 5590; 600dpi. Adobe Acrobat XI Pro was used to OCR the text and also for the merging and conversion to the final presentation PDF-format.lmchunu2014mn201

Afrotropical Culicoides : biosystematics of the imicola group. Subgenus Avaritia (Diptera : Ceratopogonidae)

A biosystematic study of seven Afrotropical and two Oriental species of the Imicola species-group was undertaken; this group of Culicoides of the subgenus Avaritia includes C. imicola the most important vector of the viruses of bluetongue (BT) and African horsesickness (AHS) known in the Old World. Five African species are redescribed i.e. C. imicola, C. pseudopallidipennis, C. bolitinos, C. miombo and C. loxodontis. Two new species are described, and the extralimital C. brevitarsis and C. nudipalpis are discussed where relevant. These nine species comprise the Imicola group, one of 10 groups constituting the subgenus worldwide. Due to confusion in the literature, the Imicola group is redefined and distinguished from the Orientalis group (also redefined); 21 species of approximately 70 world species of Avaritia are reassigned to either of the two groups. A key to all nine known species of the lmicola group is given; shortcomings in the taxonomy of the Orientalis group are discussed. The adult morphology of both sexes of the nine Imicola group species was studied; this revealed deficiencies in the descriptive format currently used in taxonomic studies of world Culicoides. Accordingly, greater detail has been introduced into descriptions and includes the use of new character states. Methods for mounting Culicoides on glass slides are also improved to ensure more accurate observation and measurement of diagnostic taxonomic features; furthermore, the descriptions are based on long series of each sex. Illustrations were made from specimens mounted symmetrically, and no feature omitted from any bodypart illustrated. Certain aspects of the life-cycle of most species were investigated but especially those of C. bolitinos, C. loxodontis and C. kwagga. The immatures of these develop exclusively in the dung of the elephant, the buffalo, the blue wildebeest, both species of rhinoceros and Burchell's zebra. Two of these species, C. bolitinos and C. kwagga, have broadened their resource range as they can invade and mature in the dung of cattle and horses. This has obvious implications for the transmission of viruses, especially where indigenous herbivores are run with domesticated livestock. In an intensive two-year survey a comparison was made between the Culicoides fauna of a natural area, the Kruger National Park (KNP), and that found in areas changed by Man, namely livestock farms adjoining the KNP. This part of the eastern Transvaal lowveld is a main focus of AHS in South Africa. Light-trapping, rearing from dung, and pootering off live hosts, revealed that some species of the Imicola group are exclusively associated with certain herbivores; these include the elephant and the zebra which are suspected or proven reservoir hosts for AHS. The results thus throw further light on the epidemiology of this disease, and also show that Man plays a decisive role in determining the numbers, and distribution, of particular Imicola group species under certain conditions. In the case of C. imicola, the commonest and most widespread of all species, this range expansion, or establishment of foci, is due to man's maintenance of domesticated livestock in confined species, and especially where these are kept on irrigated pastures. However, the serendipitous discovery of a large imicola-free zone in South Africa indicates that edaphic conditions likely play an even more important role than Man and climate in determining the prevalence and abundance of C. imicola. This area is the sandy dune field west of Port Elizabeth and holds promise as a natural quarantine zone for the import and export of livestock. AFRIKAANS : 'n Biosistematiese ondersoek van sewe Afrotropiese en twee Orientale spesies van die Imicola groep is gedoen; by hierdie groep Culicoides van die subgenus Avaritia word C. imicola wat as die mees belangrikste vektor van bloutong-(BT) en perdesiekte (AHS) virus in die Ou Wereld beskou word, ingesluit. Vyf Afrika spesies, nl. C. imicola, C. pseudopallidipennis, C. bolitinos, C. miombo en C. loxodontis, is herbeskryf. Twee nuwe spesies is beskryf asook die suid-oos Asiese spesies C. brevitarsis en C. nudipalpis word bespreek waar van toepassing. Hierdie nege spesies vorm die Imicola groep, een van die 10 groepe waaruit die subgenus Avaritia wêreldwyd bestaan. As gevolg van verwarring in die literatuur is die Imicola groep hergedefinieer en geskei van die Orientalis groep ( ook hergedefinieer); 21 spesies van ongeveer 70 wereld spesies van Avaritia is heringedeel in die twee groepe. 'n Sleutel vir al nege wereld spesies van die Imicola groep, asook 'n verspreidingskaart vir elke spesie, word gegee. Tekortkominge in die taksonomie van die Orientalis groep word ook bespreek. Die volwasse morfologie van beide geslagte van die nege Imicola groep spesies is bestudeer; dit het gebreke in die formaat wat tans vir die taksonomiese beskrywing van wêreld Culicoides gebruik word aan die lig gebring. Gevolglik is daar meer data in die beskrywings, wat nuwe karakterkenmerke insluit. Die metode van die montering van Culicoides op glasplaatjies is ook verbeter om meer akkurate ondersoek en meting van die diagnostiese kenmerke te verseker; verder, is beskrywings gebaseer op lang reekse van elke geslag. Illustrasies is gemaak van voorbeelde wat simmetries gemonteer is en geen kenmerk is uitgelaat van enige gelllustreerde liggaamsdeel. Aspekte van die lewensiklus van die meeste van die spesies, veral C. bolitinos, C. loxodontis en C. kwagga is ondersoek. Die onvolwassenes van hierdie spesies ontwikkel slegs in die mis van olifante, buffels, wildebeeste, renosters en zebras. Twee van die spesies, C. bolitinos en C. kwagga, het hulle broeimediums van voorkeur vergroot en kan eiers lê en tot volwassenheid ontwikkel in die mis van beeste en perde. Dit het vanselfsprekende gevolge vir virusoordrag tussen inheemse herbivore en vee, veral in gebiede waar die twee groepe saamloop. In 'n intensiewe twee-jaar studie, is 'n vergelyking gemaak tussen die Culicoides fauna soos aangetref in 'n ongerepte deel van Afrika, die Kruger Nasionale Park (KNP), en in gebiede aangrensend aan die KNP wat deur die mens in veeplase omskep is. Die deel van die Oos Transvaalse laeveld is bekend as 'n perdesiekte "hotspot" in Suid-Afrika. Ligvalvangste, uitbroei van Culicoides uit mis en versameling vanaf lewendige gash ere het aangedui dat sekere spesies van die Imicola groep eksklusief met sekere herbivore geassosieeris; hierby ingesluit is die olifant en zebra wat onderskeidelik verdagte en bevestigde gashere van AHS is. Die resultate dra by tot die verklaring van die epidemiologie van hierdie siekte en wys ook dat die mens 'n beslissende rol speel in die vasstelling van die getalle sowel as die verspreiding van spesifieke spesies van die Imicola groep. In die geval van C. imicola, die mees algemeenste en wydverspreidste spesie, is die uitbreiding, of daarstelling van fokuspunte, te wyte aan die mens se instandhouding van vaste bloedbanke op besproeide weiding. Die toevallige ontdekking van 'n groot imicola-vry sone elders in Suid-Afrika dui daarop dat grondtipe moontlik 'n bepalende rol kan speel in die aanwesigheid en volopheid van C. imicola. Hierdie "skoon" area is die sandduine-veld wes van Port Elizabeth en lyk belowend as 'n natuurlike kwarantyn gebied vir die invoer en uitvoer van lewende hawe.Dissertation (MSc)--University of Pretoria, 2013.Animal and Wildlife Sciencesunrestricte

Afrotropical Culicoides: C. (Avaritia) loxodontis sp. nov., a new member of the lmicola group (Diptera: Ceratopogonidae) associated with the African elephant in the Kruger National Park, South Africa

Culicoides (Avaritia) loxodontis sp. nov., is described and illustrated from both sexes collected in South Africa. It is the 5th species of the lmicola group of the subgenus Avaritia to be described from the Afrotropical Region, and is presently known only from the Kruger National Park where it has been collected in light-traps and reared from the dung of the African elephant (Loxodonta africana) on various occasions. A number of character states, and statistical analyses of antennal and palpal measurements, are used to separate the new species from its taxonomic congeners C. imicola Kieffer, 1913, C. pseudopallidipennis Clastrier, 1958, C. bolitinos Meiswinkel, 1989 and C. miombo Meiswinkel, 1991. It is suggested that the occurrence of the African elephant is the primary factor that determines the distribution of Culicoides loxodontis sp. nov., and that this close association, coupled with the fact that C. loxodontis sp. nov. can be locally abundant, may result in the cycling of certain arboviruses between this biting midge and the elephant.The articles have been scanned in colour with a HP Scanjet 5590; 600dpi. Adobe Acrobat XI Pro was used to OCR the text and also for the merging and conversion to the final presentation PDF-format.mn201

Afrotropical Culicoides : C. (Avaritia) spinifer Khamala & Kettle, 1971, a name based on an artefact (Diptera : Ceratopogonidae)

Culicoides (Avaritia) spinifer Khamala & Kettle, 1971, is made a junior subjective synonym of C. (A.) glabripennis Goetghebuer, 1935.The articles have been scanned in colour with a HP Scanjet 5590; 600dpi. Adobe Acrobat XI Pro was used to OCR the text and also for the merging and conversion to the final presentation PDF-format.lmchunu2014mn201

Discovery of a Culicoides imicola-free zone in South Africa: preliminary notes and potential significance

In December 1993, a light-trap survey was made of the Culicoides found at eight horse stables and dairies in the sandy dune field west of Port Elizabeth, South Africa. While it was notable that Culicoides numbers were low (4 749) and that the diversity was poor (15 species) , the most remarkable fact to emerge, was that C. imicola, the only proven vector of the virus of African horse sickness (AHS), was entirely absent. Though not abundant, C. bolitinos, a sister species of C. imicola, was overwhelmingly dominant (91 ,7%). Its larvae and pupae develop exclusively in the dung of cattle, but it is a species that is not implicated in the transmission of animal viruses. Elsewhere in South Africa, a frost-free climate, good rainfall and a plentiful supply of livestock would normally lead to the development of large foci of C. imicola. That this is not the case in the Port Elizabeth (P.E.) area is most likely owing to the winds inhibiting adult flight and the sandy soils being nutrient-poor and too well-drained to sustain Culicoides larvae. Studies are needed to confirm that sandy soils cannot sustain C. imicola. If so, the sandy coastal areas hold promise for quarantining against AHS.The articles have been scanned in colour with a HP Scanjet 5590; 600dpi. Adobe Acrobat X Pro was used to OCR the text and also for the merging and conversion to the final presentation PDF-format

Afrotropical Culicoides : C. (Avaritia) miombo sp. nov., a widespread species closely allied to C. (A.) imicola Kieffer, 1913 (Diptera: Ceratopogonidae)

Culicoides (Avaritia) miombo sp. nov. is described and illustrated from both sexes collected in northern Malawi. Two references in the literature have previously referred to this new species as either C. brosseti Vattier & Adam or C. imicola Kieffer. A further 4 references are discussed that most likely deal with C. miombo sp. nov. and not C. brosseti. C. miombo sp. nov. is apparently widespread in subtropical and tropical Africa and is now recorded from Zimbabwe, Botswana, South Africa, Nigeria and the Ivory Coast. There are also probable records from Angola, Burkina Faso, Zambia and eastern Madagascar. On the African mainland, both north and south of the equator, the pattern of distribution of C. miombo sp. nov. correlates strongly with that of drier Guineo-Congolian rainforest, and Sudanian and Zambezian woodlands, the latter known as miombo in southern Africa. These phytochoria and associated biota are sensitive to frost and experience relatively high temperatures and rainfall-3 factors that appear to limit the distribution of C. miombo sp. nov. to north of the 20-22°C mean annual temperature isotherms in southern Africa. The new species is a member of the Imicola group which consists of 6 species confined to the Afrotropical (including Madagascar), Oriental and eastern Palaearctic regions. One species has in historic times spread to Australia. The worldwide distribution of each species is briefly discussed. It is suggested that the Imicola and Orientalis groups are separate lineages within the subgenus Avaritia. Culicoides miombo sp. nov. is compared with its closest African congeners C. imicola, C. pseudopallidipennis Clastrier and C. bolitinos Meiswinkel; 15 character states are used to separate C. miombo sp. nov. and C. imicola. The female antennal and palpal measurements of C. miombo sp. nov. are subjected to statistical analysis to highlight their taxonomic usefulness. The larval habitat of C. miombo sp. nov. is unknown.The articles have been scanned in colour with a HP Scanjet 5590; 600dpi. Adobe Acrobat XI Pro was used to OCR the text and also for the merging and conversion to the final presentation PDF-format.mn201

Afrotropical Culicoides : a redescription of C. (Avaritia) kanagai Khamala & Kettle, 1971, reared from elephant dung in the Kruger National Park, South Africa

The discovery of Culicoides kanagai in South Africa represents a new record for this species. The female is redescribed, and the male is described for the first time. Culicoides (A.) dasyops Clastrier, 1958 is shown to be closely related to it but C. (A.) alticola is only superficially related. Short notes on the larval habitat of C. kanagai, the dung of the African elephant, Loxodonta africana, are given. The dung of both the white rhinoceros, Ceratotherium simum, and the black rhinoceros, Diceros bicornis, is considered to be a possible alternative site for the immatures of C. kanagai.The articles have been scanned in colour with a HP Scanjet 5590; 600dpi. Adobe Acrobat XI Pro was used to OCR the text and also for the merging and conversion to the final presentation PDF-format.lmchunu2014mn201

African horsesickness epidemiology : five species of Culicoides (Diptera: Ceratopogonidae) collected live behind the ears and at the dung of the African elephant in the Kruger National Park, South Africa

During the culling of elephants (Loxodonta africana) at five sites in the Kruger National Park, South Africa, a total of 682 Culicoides of five species of the subgenus Avaritia were found live either behind the ears of elephants or attracted to the freshly disembowelled intestinal dung of elephants. The species are Culicoides tororoensis Khamala & Kettle, 1971; C. kanagai Khamala & Kettle, 1971; C. loxodontis Meiswinkel, 1992, and two undescribed species, i.e. Culicoides sp. #50 and Culicoides sp. #54 pale form (p.f.). Of 511 female midges found behind ears, 39,9% were nulliparous, 57,3% empty parous, 2,5% freshly bloodfed and 0,2% gravid. The age composition of this subpopulation indicates that the Culicoides were behind the ears to suck blood and, furthermore, would do so in broad daylight. The age composition of 171 Culicoides of three species attracted to dung was entirely different: 1,8% nulliparous, 14,6% empty parous, and 83,0% gravid, indicating that the great majority of midges captured at dung were about to oviposit or had just oviposited. Immediately after culling, light-traps were operated at two of the sites. Of 4 023 Culicoides of 21 species captured, 93% were of the same five species found on the ears and at the dung of elephants. Using these and other unpublished data pertaining to the rearing of these five Avaritia species from elephant dung over the past seven years, we broadly sketch the life cycle of these Culicoides, the first for any Afrotropical species of the genus. We also discuss the implications the close association between elephant and midge has for the dispersal and geographic distribution of the latter, and how it may influence the involvement of midges in the transmission of diseases such as African horsesickness. Owing to difficulties in identifying species of the subgenus Avaritia in the Afrotropical Region, the taxonomy of each of the five above-mentioned species is briefly appraised. Of the remaining 16 species (7%) captured in light-traps 15 (6%) belong to that sector of the genus Culicoides whose immature stages develop in groundwater habitats and include C. imicola, which comprised only 2% of the light-trap collections. The large disparity in the adult abundance patterns of the "dung" and "groundwater" species in the middle of dry bushveld, is probably the result of differences in host and larval habitat preferences, and is briefly discussed. Finally, the few reports extant on the wild-host preferences of Afrotropical Culicoides are reviewed. Five tables and five figures accompany the text.The articles have been scanned in colour with a HP Scanjet 5590; 600dpi. Adobe Acrobat XI Pro was used to OCR the text and also for the merging and conversion to the final presentation PDF-format.mn201

Afrotropical Culicoides : Synhelea Kieffer, 1925, resurrected as subgenus to embrace 10 species (Diptera : Ceratopogonidae)

Synhelea Kieffer, 1925, is resurrected from synonymy to embrace 10 species of Afrotropical Culicoides. C. tropicalis Kieffer, 1913, is the subgenotype. C. vagus is made a synonym of C. dutoiti while apomorphies common to the remaining species camicasi, congolensis, dispar, moucheti, pellucidus, perettii, tauffliebi and vicinus are discussed. The broad similis group of Cornet & Chateau (1971), under which these 10 species of Synhelea were originally assembled, is not only maintained but expanded by a further 11 species. Various distributional and descriptive data pertinent to Synhelea throughout the Afrotropical region are tabulated, and keys to adult males and females presented. C. perettii and C. moucheti are new records for South Africa. C. dutoiti, C. perettii and C. tropicalis are recorded for the 1st time from Malawi while C. dutoiti and C. tropicalis are newly recorded from Botswana and Namibia respectively. Where available, notes on larval habitats are given. Synhelea is considered endemic to the Afrotropical region and is briefly differentiated from the 25 subgenera currently recognized worldwide. Numerous illustrations accompany the diagnosis.The articles have been scanned in colour with a HP Scanjet 5590; 600dpi. Adobe Acrobat XI Pro was used to OCR the text and also for the merging and conversion to the final presentation PDF-format.lmchunu2014mn201

Morphological confirmation of the separate species status of Culicoides (Avaritia) nudipalpis Delfinado, 1961 and C. (A.) imicola Kieffer, 1913 (Diptera: Ceratopogonidae)

Ten character states were used to compare five females and two males of two populations of C. nudipalpis from the Philippines and Timor in south-east Asia with 121 females and 167 males of 11 populations of C. imicola collected throughout its Afro-Asiatic range. It is concluded that they are closely related but good species, and together form a subgroup, or species-pair, in the lmicola group; currently this Old World group of the subgenus Avaritia comprises seven described species. Culicoides nudipalpis and C. imicola are most reliably separated on the ratio of the length of the proboscis to the height of the head (P/H ratio: 0,66-0,73 in nudipalpis, 0,82-1 ,02 in imicola). Compared to C. imicola very little is known of the life habits of C. nudipalpis. The latter's close taxonomic relationship to C. imicola, a proven vector of African horse sickness (AHS) and bluetongue (BT), indicates that the capacity of C. nudipalpis to vector these orbiviruses deserves to be investigated.The articles have been scanned in colour with a HP Scanjet 5590; 600dpi. Adobe Acrobat X Pro was used to OCR the text and also for the merging and conversion to the final presentation PDF-format.European Commission contract (IC 18-CT95-0010).mn201