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Culicoides truuskae Labuschagne and Meiswinkel sp. n. is described and illustrated in both sexes from material collected in South Africa and Namibia. It is restricted to the xeric western margin of the subcontinent, occurring in Fynbos, Nama-Karoo and Succulent Karoo ecoregions in South Africa and Desert and Savanna ecoregions in Namibia experiencing 600 mm of rainfall annually. Culicoides truuskae sp. n. is part of the Afrotropical 'plain-wing' Culicoides in which the wing lacks a distinguishing pattern of light and dark spots; the diagnostic dark smudge that traverses wing cell r3 may result in C. truuskae sp. n. being misidentified as the sympatric but phyletically unrelated Culicoides herero (Enderlein) - (of the Similis group, subgenus Oecacta Poey). Additionally, this study is the first description of the male of C. herero. C. truuskae sp. n. and Culicoides coarctatus Clastrier and Wirth share similar characters in the male genitalia, although the two species are separable on wing pattern and female flagellum sensilla coeloconica (SCo) distribution. The breeding habitat and adult female blood-feeding preferences of C. truuskae sp. n. are not known. A maximum likelihood phylogenetic tree, using mitochondrial cytochrome c oxidase I (COI) sequence data, is provided to further clarify the relationship between C. truuskae sp. n., C. coarctatus and C. herero. Extensive light trap data, collected over 30 years, are used to map the distribution ranges of C. truuskae sp. n., C. coarctatus and C. herero in Southern Africa.Contribution: The description of this new species and the description of the male of C. herero increases our understanding of the diversity and distribution of Culicoides species in southern Africa.

The viruses causing the economically important livestock diseases of African horse sickness (AHS) and bluetongue (BT) are transmitted by biting midges of the genus Culicoides (Diptera, Ceratopogonidae). In the Old World the most important vectors of these diseases are Culicoides imicola Kieffer, 1913, Culicoides brevitarsis Kieffer, 1917 and Culicoides bolitinos Meiswinkel, 1989. All three of these vectors belong to the Imicola complex of the subgenus Avaritia Fox, 1955. This species complex now comprises 12 sibling species; ten occur in sub-Saharan Africa and are difficult to identify (based mostly on subtle variations in the wing latterns) and so additional methods of reliable identification are needed. The pupal exuviae of the five commonest sibling species (C. imicola, C. bolitinos, Culicoides loxodontis Meiswinkel, 1992, Culicoides tuttifrutti Meiswinkel, Cornet & Dyce, 2003 and Culicoides sp. # 107) harvested from a variety of large herbivore dung types and from decaying fruits, are described and illustrated in detail. It is shown that they can be differentiated clearly on a number of morphological characters and, furthermore, are separable into two distinct groups based (principally) on the shape of the respiratory organ. A key for identifying and differentiating these five pupae is provided. Also, the pupa of the Oriental-Australasian C. brevitarsis was compared with its allopatric sister taxon, C. bolitinos. Because they share a common larval habitat (cattle and buffalo dung) and are almost inseparable in the adult phenotype, the question of their possible synonymy is raised. However, their respective pupae could not be differentiated on gross morphology and so it is argued that this unresolved problem requires a molecular solution.

In December 1995 to March 1996 and the early summer of 1997 South Africa experienced above average rainfall which favoured the occurrence of Culicoides transmitted diseases. During this period several outbreaks of an uncommon disease of cattle occurred over a large part of the country. The clinical signs were similar to those of infection with the viruses of bluetongue (BT) and epizootic haemorrhagic disease of deer (EHD). Virus isolation from cattle and Culicoides yielded both viruses. Dual infections occurred on several farms. Typing of BT isolates yielded types 2, 3, 6 and 8. On at least two farms more than one BT virus serotype was involved. On one farm only EHD virus could be isolated from cattle and Culicoides. Serological tests confirmed that on this farm the disease was caused by EHD. In 1932/33, when a similar disease was reported conditions were vastly different. Rainfall figures show that the 1932/33 season was exceptionally dry. Techniques available at that time could not identify EHD and the cause was reported to be BT. The occurrence of BT in a dry season and over a much wider area than the distribution in South Africa of Culicoides imicola, the only proven vector for BT, is a clear indication that other species less dependent on high rainfall are involved. The present isolation of BT virus from three of five pools of parous C. bolitinos is evidence that this species, which breeds in cattle dung, may be an additional vector for BT.

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 imicola 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.

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 (4749) 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.

In 54 light-trap collections made at 28 sites in the Onderstepoort area a total of 178,941 Culicoides midges of 35 species was collected in March 1988; the survey was repeated at 26 sites in September and yielded 19,518 Culicoides of 24 species. The number of Culicoides species collected totalled 38. C,imicola was the most abundant species at 27 of the 28 sites sampled, and accounted for 88% and 67% of all midges collected in the two months respectively. This study not only confirms that C. imicola is widespread and abundant in the greater Onderstepoort area, but also that its numbers correlate positively with the historical prevalence of African horse sickness (AHS) and bluetongue (BT) locally. The high numbers of C. imicola make Onderstepoort the ideal site for the study of its laboratory vector capacity. The relatively low numbers of Culicoides spp. other than C. imicola in the Onderstepoort area, will severely limit studies on their roles in the transmission of arboviruses. The origin of the blood-meals of 1338 engorged Culicoides belonging to 13 species was determined by means of cross-over electrophoresis precipitin test; C. imicola fed on cattle, horses, sheep of pigs, Four other Culicoides species showed a similarly wide host range.

Altogether 52 078 Culicoides biting midges of 35 species were collected during February 1990 and 1993 in 40 light-trap collections made on 17 cattle and/or sheep farms in the Bethlehem and Fouriesburg districts of the colder, high-lying eastern Orange Free State. Culicoides (Avaritia) bolitinos was by far the most abundant species, representing 50.9% of all specimens collected. Culicoides (A.) imicola, considered to be the most common stock-associated species in the summer rainfall areas of southern Africa, and the only proven vector of bluetongue virus (BTV) and African horsesickness virus (AHSV) in the subregion, was uncommon, comprising only 1.4%. While AHS is apparently absent, BT and bovine ephemeral fever (BEF) are endemic in this cooler, high-lying area of South Africa. The virtual absence of C. imicola implies that other Culicoides species, such as C. bolitinos and C. cornutus, may be involved in transmitting BT virus (and perhaps BEF) in the eastern Orange Free State, and possibly elsewhere in Africa. Virus isolation attempts made on 45 single species pools of C. bolitinos, C. pycnostictus, C. milnei, C. leucostictus, C. zuluensis and C. gulbenkiani were, however, negative. Finally, 20 of 28 blood-engorged Culicoides of 11 species, which were tested against cattle, sheep, horse, pig and bird antisera, tested only positive against cattle antisera.

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.

Analyses of 2 light trap catches comprising 6,041 and 1,598 Culicoides showed that the reliability of subsampling such catches increased with subsample size, while the subsampling error decreased with an increase in the number of individuals per species present in a subsample. Subsamples comprising approximately 500 Culicoides are deemed sufficient for comparing population densities between sites and sampling occasions and also give an acceptable indication of relative species abundance at a site. It is recommended that species for which the mean number of specimens in subsamples originating from 3 catches at a site is less than 7, should not be included in biometric analyses aimed at comparisons of population densities. For all other species a test level of 1% rather than 5% should be used for such comparisons. When species ratios obtained from subsampled catches are employed as indicators of abundance, the chi-squared test should be utilized at a 1% level if the ratios originate from 3 catches at a site and at a 0.1% level if only one catch per site is made. Due to poor representativeness of small catches, it is suggested that species for which fewer than 7 individuals are present in a single subsample, be excluded from chi-squared tests. A 5-point procedure for subsampling a large light trap collection of Culicoides is given.

Culicoides (Avaritia) loxodontis sp. nov., is described and illustrated from both sexes collected in South Africa. It is the 5th species of the Imicola 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.

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 degrees 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.

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.

A total of 33,564 Culicoides midges was collected in 44 light trap collections made at 22 sites in the Stellenbosch area during November 1986. Of the 23 species present in these collections 8 were frequently encountered namely, C. magnus, C. imicola, Culicoides sp. 49, C. zuluensis, C. gulbenkiani, C. pycnostictus, C. distinctipennis and C. nivosus. Although C. magnus was abundant at all trap sites, the prevalence of the other species appeared to be affected by the proximity of the light trap to different host animals and/or larval habitats. Plain-wing species and members of the C. schultzei group were rarely collected. The larval habitats of most of the above species were located by the use of tent-type emergence traps. All these habitats were found on irrigated pastures or where drainage water had accumulated. The difference in the requirements of the various species was associated with certain factors, such as degree of moisture, the type and amount of organic matter present and the particle size of the underlying soil. The identity of the blood-meals of 69 individual Culicoides belonging to 7 species was determined. The 5 commonest species had all fed on cattle and 4 of these on sheep. Two species, C. pycnostictus and C. distinctipennis were positive for bird blood.

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.

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.