"8. |OiS" 26AUG1982 Bulletin of the British Museum (Natural History) Entomology series Vol 44 1982 British Museum (Natural History) London 1982 Dates of publication of the parts No 1 . .28 January 1982 No 2 25 February 1982 No 3 25 March 1982 No 4 . . 29 April 1982 ISSN 0524-6431 Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset Contents Entomology Volume 44 Page No 1 The taxonomy, biology and medical importance of Simulium amazonicum Goeldi (Diptera: Simuliidae), with a review of related species. A. J. Shelley, R. R. Finger & M. A. P. Moraes 1 No 2 A revision of the genus Belonogaster de Saussure (Hymenoptera : Vespidae). O. W. Richards . 31 No 3 The taxonomy and phylogeny of the genus Polyura Billberg (Lepidop- tera: Nymphalidae). Robert L. Smiles . 115 No 4 A taxonomic revision of the genus Gastrimargus Saussure (Orthoptera : Acrididae). J. Mark Ritchie 239 T» 11 *.• rxi_ ' Bulletin of the *** _ British Museum (Natural History) The taxonomy, biology and medical importance of Simulium amazonicum Goeldi (Diptera : Simuliidae), with a review of related species A. J. Shelley, R. R. Finger & M. A. P. Moraes Entomology series Vol 44 No 1 28 January 1 982 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England. World List abbreviation : Bull. Br. Mus. nat. Hist. (Ent.) Trustees of the British Museum (Natural History), 1982 ISSN 0524-643 1 Entomology series Vol 44 No 1 pp 1-29 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 28 January 1982 -* 78 77 Figs 74-79 S. amazonicum pupa. 74, frontoclypeus ; 75, thorax ; 76, platelets and spinules of thorax ; 77, abdomen, dorsal view; 78, abdomen, ventral view; 79, spiny cuticular processes of eighth sternite. mium : distance between apex of corner tooth and first hypostomial seta slightly less than that between corner teeth. Antenna long (Fig. 84), unpigmented; first segment about five times as long as broad, segment length ratios about 5 : 4.8 : 5.5 (only one antenna examined). Mandible (Fig. 85) with one narrow elongate mandibular serration, sometimes with small serration posterior to this (Fig. 86); first three comb teeth equal in size; ventral margin of mandible posterior to mandibular serration with row of saw-like serrations (Fig. 87). Maxillary palp short, about three times as long as breadth at base (Fig. 88). Cephalic fan with 12- 15 rays. SIMULIUM AMAZONICUM GOELDI 17 86 88 Figs 80-92 S. amazonicum larva. 80, larva, dorsal view; 81, head, dorsal view; 82, head, ventral view (flattened); 83, hypostomium ; 84, antenna; 85, mandible; 86, mandibular serrations; 87, mandible, ventral margin snowing saw-like serrations; 88, maxillary palp; 89, cuticular spine; 90, lateral sclerite of proleg; 91, pupal respiratory histoblast; 92, posterior end of abdomen, lateral view, showing ventral papillae. 18 A. J. SHELLEY, R. R. FINGER & M. A. P. MORAES 93 Figs 93, 94 S. amazonicwn larva. 93, anal gills ; 94, anal sclerite. Thorax yellow to light brown with scattered dark mottling of variable form dorsally and one or two central patches posterior to proleg ventrally. Cuticle of dorsal surface densely covered with fine spines (Fig. 89), lateral surface with scattered spines and ventral surface bare. Proleg circlet with about 22 rows of 1-6 hooks, lateral sclerite faintly pigmented with about eight processes (Fig. 90). Pupal respiratory his- toblast (Fig. 91) with eight tightly coiled filaments divided into three primary branches near base. Abdomen yellow to brown with grey annular mottling on four anterior segments and varying patterns on other segments. Cuticle with spines arranged as on thorax except scattered spines present ventrally also. Ventral papillae small (Fig. 92). Scattered short spines on anterior fold of anus, tri-lobed rectal gills with about eight secondary lobules on each lobe (Fig. 93). Anal sclerite (Fig. 94) with arms strongly sclerotised; posterior arm extending to between ninth and twelfth rows of posterior circlet hooks. Posterior circlet with about 54-56 rows of 2-12 hooks. 2 + 2 (sometimes 1 + 1) mid-dorsal and 1 + 1 lateral strongly developed setae between posterior arms of anal sclerite and posterior circlet. MATERIAL EXAMINED Syntypes $, Brazil: Amazonas, Bom Lugar, R. Purus, v.1904 (Huber) (25 9 in alcohol, pinned or as slide preparations, BMNH; 27 9, pinned, NM). [The International Commission on Zoological Nomenclature has been requested to set aside these syntypes under their plenary powers (Shelley, 1981).] Brazil: 1 9, Amazonas, Valparaiso (near Lagoa Bom Lugar), R. Purus, approx. 8°42'S 67°22'W, approx. elevation 120 m, ex pupa, 22.xi.1977 (Shelley) (BMNH). [The International Commission on Zoological Nomenclature has been requested to designate this specimen as a neotype under their plenary powers (Shelley, 1981).] Topotypes. 18 9, ex pupae (pinned and as slide preparations); 31 <£, ex pupae (pinned and as slide preparations); 16 pupae (in alcohol); 10 larvae (slide preparations); data as for proposed neotype. Numerous 9, collected from human bait and horse (pinned, in alcohol, slide preparations); data as for proposed neotype. (AMNH, BMNH, IOC, MLP, USNM.) Numerous 9, collected from human bait (pinned, in alcohol); locality data as for proposed neotype, 29.xi.-3.xii. 1976, 17-19.vi.1977 (Arouck) (BMNH, IOC). IJ, Amazonas, Igarape Escondido (Lagoa Bom Lugar), R. Purus, ex pupa, 17.xi.1977 (Shelley) (BMNH). DISTRIBUTION. Because of the previously confused situation over the identity of 5. amazonicum the confirmed distribution of this species is based only on material studied by the authors for the present work. The distribution of this species in relation to other species in the S. amazonicum- group that occur in Brazil is shown in Fig. 95. S. amazonicum was found at the following localities in addition to the type-locality on the Purus river system, the specimens having been deposited in the BMNH and IOC: several 9, Boca de Acre, confluence of R. Acre and R. Purus, 8°45'S 67°24'W, 24.xi.1977 (Shelley); numerous 9, Labrea, R. Purus, 7°14'S 64°50'W, 8.X.1973 (Shelley); numerous 9, Capacini, near Labrea, R. Purus, 25. ix. 1976 (Shelley); numerous 9, Parana do Ituxi, confluence of R. Ituxi with R. Purus, near Labrea, 30.ix.1976 (Shelley). Specimens tentatively determined as S. amazonicum have also been collected biting man at the following localities (slight scutal variations are apparent from the Bom Lugar form but their significance cannot be decided until immature stages and males have been examined). Numerous 9, Amazonas State, Feijoal, R. Solimoes, 4°10'S 69°25'W, 5.x. 1979 (Shelley & Luna Dias); numerous 9, Tefe, R. Solimoes, 3°22'S 64°43'W, 8.x. 1978 (Shelley & Luna Dias); numerous 9, Roraima Territory, Km 50, Northern Perimeter Road (BR 210), R. Ajarani, 1, 2°01'N 61°28'W, 16.U979 (Shelley & Luna Dias). SIMULIUM AMAZONICUM GOELDI r A S.»m«ionicum A S. mlnuaculum 0 S.quidr if idum 1 S.roraim«n»« VSimuhum ap.(Mad»ira) / 200km Fig. 95 The distribution of species of the S. amazonicum-group in Brazil. Inset : detail of Roraima Territory. (Based on reared specimens and type-material.) BIOLOGY. Although various aspects of the biology of S. amazonicum have already been investi- gated by several workers in Brazil and neighbouring countries their value is dubious as it is probable that other species were involved. Studies on the biology of this species as well as other anthropophilic species in the S. amazonicum-group that are large-river breeders are hampered by the difficulties involved in sampling for immature stages. Breeding grounds, even in localities where adults attack man in enormous numbers, are difficult to find. Apart from the more obvious difficulties involved in sampling large rivers, such as water depth, current speed and the time and manpower needed to examine, vast areas of substrate for larvae and pupae, account must be taken of the large fluctuation in river depth over a short period of time principally in the rainy season. Even large rivers in the Amazon basin may rise by as much as a metre over night as a result of rainfall often some considerable distance away. Detailed investigations at Valparaiso, near Bom Lugar, the type-locality of S. amazonicum, were not possible and hence only brief comments are made mainly on larval and pupal habitat. However a study made some years ago at Capacini, also on the R. Purus, by one of the authors (A.J.S.) is cited, now that the correct identity of 5. amazonicum has been established. 20 A. J. SHELLEY, R. R. FINGER & M. A. P. MORAES Studies at Valparaiso. The river Purus at Valparaiso is about 100 m wide, has a fast current and few suitable objects for mooring a boat to facilitate sampling. The banks of the river, with its many meanders, are either steep and devoid of vegetation, in the form of sandy spits, or marshy and covered with herbaceous vegetation and scattered bushes. Immature stages of S. amazonicum were found only on a small submerged wild guava bush near the river bank in strong current up to 1.5 m below the water surface. This was the only breeding ground found after four days of searching in the river Purus at a time of the year when swarms of S. amazonicum attack man at the river's edge throughout the day. Pupae were found mainly on small branchlets but sometimes attached to leaves, and were usually covered in sediment. No pupae or mature larvae of other Simulium species were found. Collections from several forest streams draining into the R. Purus or into Lagoa Bom Lugar produced large numbers of immature stages of 5. quadrifidum but only one male S. amazonicum pupa. In a collection from human bait several thousand S. amazonicum females were identified as well as one female Simulium sp. (Madeira), which has been shown to be a vector of M. ozzardi on the R. Solimoes (Shelley et a/., 1980). 5. amazonicum was also collected feeding on a horse at Valparaiso. Studies at Capacini. Aspects of the biology of S. amazonicum were studied at Capacini, a small rubber collecting community near Labrea on the R. Purus, in low lying (altitude 60 m) tropical rain forest. The observations were made during investigations into the vector of M. ozzardi at this locality (Shelley & Shelley, 1976) but not hitherto published because of uncertainty as to the identity of the species. Reexamination of voucher specimens (deposited in the BMNH ahd IOC) has confirmed that true S. amazonicum is the species involved. r. . ,• . ... Only S. quadrifidum was found in the two local streams, Igarape Capacini and Igarape -Ca-Te- Espera, and no immature stages were found in the R. Purus. Studies-^ere therefore confined to adult S. amazonicum, the only anthropophilic black-fly found in the area.Xv. • To discover the resting sites of 5. amazonicum searches with a sweep net wete-made in potential resting sites alongside the river and at the forest's edge by the two streams. Host preference was determined by making a 12-hour catc|x'-using.bak?of man, C9W, sheep, dog, monkey and chicken. These were positioned, 10 m apart, in bean' fields at the 'edge of -the river and three collectors captured all flies biting from 06.00 (o .1,8.00 hrs; two. further collectors captured flies biting a human and a bovine bait in the cattle pastures above the bean fields. To determine the biting pattern of S. amazonicum five collections were made from human volunteers clad in shorts and seated in the bean fields for three consecutive days. One collector was assigned to each bait and commenced capturing flies at 06.00 hrs (dawn) until midday; he was replaced by another col- lector who worked from midday to 18.30 hrs (dusk). Flies were collected directly into alcohol, a small number being set aside for taxonomic studies, the rest being dissected later for filariae. Daily records of temperature and relative humidity were kept. No S. amazonicum were found on vegetation at the forest edge but 29 females were collected from long grass in the cattle pasture and 15 females from cracks in the soil in the bean fields. S. amazonicum showed a marked preference for human blood although the cow could be an import- ant secondary host in the area (Table 1). During the three consecutive days of the biting catch Table 1 Biting preferences of S. amazonicum at Capacini Bait Bean fields Pasture No. collected % of total catch No. collected % of total catch Man 794 90.7 1292 80.1 Cow 80 9.1 321 19.9 Sheep 1 0.1 0 — Dog 0 — 0 — Monkey 0 — 0 — Chicken 0 — 0 — Totals 875 99.9 1613 100.0 SIMULIUM AMAZONICUM GOELDI 21 35- 33i 31 29- 27- /**- ^ ^^^. TEMPERATURE ^^ X RELATIVE ^*^ ' HUMIDITY 80 -70 HOURS LOCAL TIME Fig. 96 The biting cycle of S. amazonicum at Capacini. 3 400 S. amazonicum were collected from the human baits. The biting pattern is shown in Fig. 96 where the geometric mean hourly catch (Mw , after Williams) expressed as a percentage of the total days' catch (% Mw) is plotted against time. Biting occurred between dawn and dusk with the main peaks at 07.00-08.00 and 15.00-18.00 hrs local time. MEDICAL IMPORTANCE. Formerly S. amazonicum was considered to be a noxious man-biting species in several countries in South America and many published reports concerning this exist in the literature. It is impossible to establish which, if any, of these reports actually refer to S. amazonicum as specimens were not deposited in collections at the time. However, investigations have been made where this species has been cited as a vector of pathogens to man. Only two have been found, both involving human filariae. In Brazil Cerqueira (1959) named S. amazonicum as the vector of M. ozzardi at Codajas on the R. Solimoes. No critical analysis of Cerqueira's data had been made until Shelley et al. (1980) challenged his conclusions on the data that he had presented, as well as on his identification of the vector. Specimens from Codajas identified by Cerqueira as S. amazonicum have now been determined (A.J.S.) as an undescribed species of the S. amazonicum-gToup referred to in this paper as Simulium sp. (Madeira) (p. 27). That this species is a vector of M. ozzardi in Brazil has now been confirmed by these authors (Shelley et al., 1980) at Feijoal on the river Solimoes. They also showed that 5. amazonicum, a less common species at this locality, is also a vector of this filaria. This species is responsible for transmission of man- sonelliasis along the river Purus as Shelley & Shelley (1976) found a female S. amazonicum naturally infected with M. ozzardi at Capacini, and more recently an infective larva of this filaria was recovered from an experimentally infected fly that had previously fed on a person with mansonelliasis at Valparaiso (A.J.S., unpublished data). The supposed incrimination of 5. amazo- nicum as a vector of onchocerciasis at the R. Toototobi in Brazil by Rassi et al. (1975) is an error stemming from their misidentification of 5. minusculum as 5. amazonicum. A summary of the distribution, host preference and vector capacity of species in the S. amazonicum-group is given in Table 2. 22 A. J. SHELLEY, R. R. FINGER & M. A. P. MORAES Table 2 The distribution, host preference and vector capacity of species in the S. amazonicum- group (based on a report by the World Health Organization, 1979). Species Distribution Host preference Vector of Man Animal M. ozzardi 0. volvulus S. amazonicum Brazil + + + ? Colombia + + S. chaquense S. minusculum Argentina Brazil ?+ - - + Colombia + + — Guyana Venezuela + - - + S. quadrifidum S. sanguineum Simulium sp. (Barbacoas) Simulium sp. (Madeira) Brazil Colombia Venezuela Brazil Colombia t 7 - : Taxonomic notes on other species in the 5. amazonicum-group Simulium chaquense Coscaron Simulium chaquense Coscaron, 1971: 33. Holotype 9, ARGENTINA: Chaco, Arroyo Zapirain, approx. 27°S 59°W, ex pupa (MLP) [examined]. [Coscaron (1971) gives the type-locality as Formosa Prov- ince. Arroyo Zapirain is a stream running through a small farm crossed by Route 1 1 on the border of Chaco and Formosa Provinces; the actual collection site is in Chaco Province (Coscaron, pers. comm.).] This is a recently described species only known from one locality in northern Argentina. It has most of the characters of the S. amazonicum-group but is exceptional in having two distal spines on the male distimere whereas all other species in the group possess one. The extent to which S. chaquense feeds on man is uncertain but it does not appear to be of medical importance. MATERIAL EXAMINED Simulium chaquense Coscaron, holotype 9 (ex pupa), Argentina: Chaco, Arroyo Zapirain, 14.vii.1971 (Coscaron) (MLP). Argentina: 5 9, 2 <3 (ex pupae), data as for holotype (2 9, 1 (J in BMNH;3 9, 1 3 in MLP); 1 9 (ex pupa), Corrientes, Arroyo Santa Maria, 9.vii.l971 (Coscaron) (MLP) (all paratypes of Simulium chaquense Coscaron). Simulium minusculum Lutz Simulium minusculum Lutz, 1910: 253. Syntypes 9, BRAZIL: Minas Gerais, Lassance, Rio das Velhas (IOC) [examined]. [Synonymised with S. amazonicum Goeldi by Lutz, 1917: 64; resurrected from synonymy by Cerqueira & Nunes de Mello, 1964: 102.] Simulium miniusculum: Malloch, 1912: 653. [Incorrect subsequent spelling.] [Simulium amazonicum Goeldi sensu Lutz, 1917: 63 and subsequent authors except Smart, 1942: 46 and Cerqueira & Nunes de Mello, 1964: 102(9 only). Misidentifications.] ? Simulium roraimense Nunes de Mello, 1974: 45. Holotype <$, BRAZIL: Roraima, Cachoeira, R. Cauame, ex pupa (INPA) [not examined, unlocated in type-depository]. S. minusculum is the common Brazilian species of lowland tropical forest that has been most often confused with S. amazonicum. The three most significant contributors to the morphology and medical importance of S. minusculum (as S. amazonicum) have been Lutz, Porto and Rassi. Each is dealt with separately. SIMULIUM AMAZONICUM GOELDI 23 LUTZ. The major Brazilian contribution to the morphology of the species that we regard as S. minusculum in this paper was made by Lutz who believed that he was dealing with S. amazonicum. Because of the rather involved nature of Lutz's taxonomic studies on S. minusculum and allied species, a summary of his work covered by his 1909, 1910 and 1917 papers is given followed by our interpretation of his findings. His first reference (Lutz, 1909) to 5. amazonicum simply quoted Goeldi's original description (1905) and described a new species, S. exiguum (Lutz, not Rouband) from females collected at the R. Grande, near Franca in Sao Paulo state. This was followed in 1910 by a description of a new species, S. minusculum, from females sent in alcohol by Chagas from Lassance, Minas Gerais. In the same paper he compared these with specimens, also sent in alcohol, from the Madeira-Mamore region in Rondonia and judged them to be conspecific, but at the same time noting differences in the pruinose pattern of the female scutum due, possibly, to different storage times in alcohol of the material from these two localities. He also noted that, apart from leg coloration, S. minusculum closely resembles S. amazonicum, and suggested their possible synony- my. Pupae with four gill filaments, collected from a stream near the Madeira-Mamore railroad and assumed to be S. minusculum, were described. Having examined more specimens of 'S. amazonicum ' from localities in the Territory of Rondonia, states of Bahia and Minas Gerais, as well as the river Tocantins (state not indicated), Lutz (1917) observed that the angle of incident light affects the form of the pruinose scutal pattern and that this pattern is altered by alcohol preservation and hence is only clear in fresh or dried material. He also noted that darkening of tissues, particularly in the legs, occurs following a blood meal. He therefore attributed the differences previously noted between S. minusculum and S. amazonicum to these causes, sank the former species as a synonym of S. amazonicum and suggested the possible synonymy of S. exiguum (Lutz, not Roubaud) with this species. Furthermore, he sank S. nitidum Malloch, from Huancabamba, Peru, as a synonym of S. amazonicum after examining specimens but not the 'type' from this locality. As his previous description of '5. amazonicum' in 1910 was based on spirit material, Lutz (1917) redescribed both the female and pupa of 'S. amazonicum'' from fresh material obtained by mass rearing six-filamented pupae collected at Sant'Anna de Sobradinho on the R. Sao Francisco above Joazeiro, Bahia. The four-filamented pupa collected at Madeira-Mamore and previously attributed (Lutz, 1910) to 5. amazonicum was given the name S. quadrifidum. The synonymy (Lutz, 1917) of S. minusculum with S. amazonicum cannot be upheld. Apart from coloration Lutz's main criterion for erecting S. minusculum in 1910 was its small size (body length 1.00-1.25 mm). Thirteen of Lutz's original series of pinned syntypes from Lassance, Minas Gerais (nos. 12552-12555, 12558) have been examined and the body length (eight specimens measured) found to range from 1.4-2.1 mm; the scutal pattern, only discernible in the. better preserved specimens, is distinct from that of S. amazonicum but similar to that of the S. sanguineum holotype, with which the specimens were compared. Three of Lutz's slides were also examined : no. 12001 with two six-filamented pupae from Madeira-Mamore, labelled as S. amazonicum but not referred to in any of his papers, no. 12005 with a male and pupal pelt, and no. 12007 with eight six-filamented pupae from Sant'Anna do Sobradinho labelled as S. amazonense (error for S. amazonicum used by Lutz, 1917), which in 1917 he considered conspecific with the S. minusculum specimens from Lassance. In their redescription of S. amazonicum Cerqueira & Nunes de Mello (1964) referred to Lutz's material and revalidated S. minusculum for the specimens reared from six-filamented pupae collected at Sant'Anna do Sobradinho as well as the females from the type-locality at Lassance, but considered the females collected at Madeira-Mamore as S. amazon- icum. Until reared material is available from Lassance it is convenient to accept Cerqueira & Nunes de Mello's revalidation of S. minusculum for Lutz's material collected from the first two localities as well as the females and six-filamented pupae in the Lutz collection from Madeira-Mamore. Cerqueira & Nunes de Mello regarded material from the last locality as S. amazonicum but they had confused three species of the S. amazonicum-group in their 1964 redescription. Our collections of six-filamented pupae from the R. Jacy Parana in the Madeira-Mamore region gave rise to females indistinguishable from Lutz's S. minusculum from Lassance. As S. sanguineum s.str., which at present is only distinguishable from S. minusculum by 24 A. J. SHELLEY, R. R. FINGER & M. A. P. MORAES its eight-filamented pupa, has not been recorded in Brazil, females from Brazil without associated pupal pelts that key out to S. sanguineum or S. minusculum are provisionally regarded as S. minusculum. S. chaquense and S. roraimense, also inseparable from S. minusculum in the female, are also excluded due to the restricted distribution and the uncertainty surrounding the validity of the species respectively. The main vector of 0. volvulus at Toototobi, tentatively determined as S. sanguineum (Shelley et al., 1979) and subsequently, when the pupa of 5. sanguineum became known, as S. sanguineum s.l. (Shelley et al., 1980) until it could be assigned to a species, now falls provisionally under S. minusculum. PORTO. Porto (1939) accepted Lutz's synonymy (1917) of S. minusculum with 5. amazonicum as he believed the morphological variations described by Lutz in his material not to be interspecific. Although Porto (1939) did not see type-material of S. amazonicum he redescribed this species from females collected in the states of Mato Grosso and Bahia, which are now in the Public Health Faculty of Sao Paulo University (see Forattini et al., 1971). Vulcano examined them as well as additional specimens from other states in 1943, and concluded that all were two infra- specific taxa of S. amazonicum which she labelled with (unpublished) manuscript varietal names. The senior author examined this material and recognized three species among the pinned speci- mens, none of which is 5. amazonicum: specimen nos. 4345-4348 and 4350-4352 from Rio das Mortes, Mato Grosso as well as no. 1 157 from Rio de Cobre, Salvador, Bahia were provisionally determined as S. minusculum; specimen nos. 4330-4340 from Rio das Mortes, Mato Grosso are of an undetermined species. The additional material determined as S. amazonicum by Vulcano and also included in the Forattini et al. catalogue (1971) are as follows: specimen nos. 4554-4555 from Acre state are now provisionally determined as S. minusculum; specimen nos. 4309-4312 from Porto Cabral, Sao Paulo, 4313^316 from Rio Claro, Sao Paulo, 4321-4323 from Dou- rados, Mato Grosso and 4324-4326 from Juquia, Sao Paulo and Itatiaia and Teresopolis, Rio de Janeiro are now determined as 5. incrustatum. RASSI. Rassi's two publications, resulting from a visit in 1974 to the Brazilian onchocerciasis foci, are important as they refer to S. amazonicum as a vector of 0. volvulus in Brazil. The flimsy evidence provided by Rassi et al. (1975) for this incrimination has been discussed elsewhere (Shelley et al., 1979). In his earlier paper Rassi (1974) refers to S. amazonicum as a vector but acknowledges the difficulties involved in the identification of this species, which he considers to be a complex. He recognizes two species of this ' complex ', which he separates on female size, to be widespread in the Brazilian Amazon, including the onchocerciasis foci. The commoner and smaller species he refers to as S. amazonicum Goeldi Lutz (1910, 1917), which he maintains corresponds to S. minusculum and S. quadrifidum as described by Lutz, and the larger as S. amazonicum Goeldi as described by its original author. A redescription of the smaller species is given and it is this species that Rassi (1974) and Rassi et al. (1975) believe to be the vector of 0. volvulus in Brazil. In this redescription based on material collected at Toototobi, females were found to have a body length of 1.0-1.5 mm compared to Goeldi's reference to 2.06 mm body length for S. amazonicum. Pupae with four gill filaments, assumed to be of the smaller species, were found by Rassi (1974) in the Toototobi river. He also assumes that the larger species, of which no body length measurements or description are given, has a six-filamented pupa, presum- ably based on Lutz's description (1917) of 'S. amazonicum'. We made collections of man-biting black-flies as well as larvae and pupae from the river Toototobi and surrounding forest streams on eight occasions over three years in both wet and dry seasons, but were unable to find the two species of the ' S. amazonicum complex ' detailed by Rassi (1974). Only one anthropophilic species of the S. amazonicum-group, now provisionally assigned to S. minusculum until the status of S. roraimense is confirmed, was found by us at this locality where it was shown to be a vector of onchocerciasis (Shelley et al., 1979). This was the most common black-fly in the area and showed considerable variation in female body length in February 1976: 1.2-2.0 mm for dried females (101 specimens measured) and 1.7-2.7 mm for females preserved in alcohol (210 specimens measured) with a normal distribution for body length. A zoophilic species, S. quadrifidum, was reared from four-filamented pupae collected in the SIMULIUM AMAZONICUM GOELDI 25 river Toototobi. It is apparent that Rassi (1974) misidentified both S. minusculum and S. quadri- fidum as S. amazonicum. The vector of 0. volvulus in the Upper Orinoco region of Venezuela, adjacent to the Toototobi in Brazil, indicated as S. amazonicum by Rassi et al. (1977) has probably been similarly misidentified. Variations in the scutal pattern of female S. minusculum collected by us from different localities in Amazonia as well as in Lutz's syntype-series from Minas Gerais have been noted. Due to the paucity of reared material from these localities and the total lack of such material from the type-locality, it cannot at this stage be decided whether S. minusculum is a polymorphic species or whether more than one species is being assigned to this name. A further complication is seen in the case of S. roraimense. Although we have examined reared topotypes the only means of separating this species from S. minusculum is by the form of the male scutal pattern. The validity of the presence or absence of merging of the three scutal vittae as an interspecific character for separating these two species is not known. As this is the only apparent difference and since S. sanguineum males show variation in the extent of merging of these vittae (Tidwell et al, 1981), S. roraimense is tentatively treated as a synonym of S. minusculum in this paper. This may be confirmed only when reared S. minusculum topotypes are available and more reared material has been collected to examine the degree of variation in this character. MATERIAL EXAMINED Simulium minusculum Lutz, 16 9 syntypes, Brazil: Minas Gerais, Lassance, Rio das Velhas. 17°53'S 44°34'W 3.i\.l91Q(Chagas) (IOC, nos. 12552, 12554 & 12558) [labelled by Lutz as S. minusculum']. Brazil: 29, 1 cJ, 1 c? (ex pupa), 8 pupae, Bahia, Sant'Anna do Sobradinho, R. Sao Fransisco, 9°23'S 40°50'W, 25.vi.1912 (Lutz) (IOC, nos. 12005, 12007, 12461, 12462) [labelled by Lutz as S. amazoni- cum, S. amazonense or S. minusculum (amazonicum)] ; 2 pupae, Rondonia, Madeira-Mamore region, R. Madeira. 1910 (Cruz) (IOC, no. 12001) [labelled by Lutz as S. amazonicum']; 5 9, 5 ^ (ex pupae), Madeira-Mamore region, Jacy Parana, R. Jacy Parana, 9°15'S 64°24'W, 16.x. 1978 (Shelley & Luna bias) (BMNH & IOC); 34 9, 34 $ (ex pupae), Roraima, Cachoeira Bern Querer, R. Branco, 1°58'N 61°00'W, 16.U979 (Shelley & Luna Dias) (BMNH & IOC); 4 <$ (ex pupae), Northern Perimeter Road (BR 210), R. Ajarani 1, 2°01'N 61°28'W, 16.U979 (Shelley & Luna Dias) (BMNH & IOC); 1 9 (ex pupa), 8 9, 8 a Ritsema) 44 Bristles of mesoscutum and humeri longer and finer, hair-like. Clypeus rarely with such coarse reticulation. Gena narrower. Mesoscutum with sparser, whitish or dense greyish tomentum 45 44(43) Wings uniformly pale greyish brown. Femora reddish. Senegal, Liberia, Ghana, Sierra Leone, Nigeria, Fernando Po, Gabon, Congo, Rwanda . B. brunnea brunnea Ritsema (p. 64) Wings dark red-brown, tips darker. Femora usually black. Uganda, Kenya B. brunnea nigriclava subsp. n. (p. 66) 45(43) Larger, wing-length 19-0 mm. Hind femora with a few pale or more rarely short black bristles beneath. Black, head except frons, usually front of pronotum, scutellum and metanotum, gastral tergites 1-2, sometimes 5-6 (more or less), red; tergite 2 sometimes with two yellow spots. Wings yellow-brown, tips dark. Stalk of second gastral tergite twice as long as broad. Propodeum with dense black hairs and usually strong, oblique striae but sometimes striae weaker and large, shallow punctures more obvious. Gaster sometimes with a few protrud- ing black bristles. Ethiopia . B. meneliki Gribodo (p. 52) Smaller, wing-length 16-0-17-5 mm. Hind femora with numerous short black bristles beneath 46 46(45) Wings brownish hyaline, length 17-5 mm. Stalk of second gastral tergite 3-5 times as long as broad. Mesosoma and gaster ferruginous, segment 4 a little darkened, 5-6 usually re- ddish, tergite 2 with a pair of transverse yellow spots. Mesoscutum without tomentum. Principe I B. principals sp. n. (p. 82) Wings brown, tips darker, length 16-0 mm. Stalk of second gastral tergite 2-5 times as long as broad. Black, head except frons in part, tibiae and tarsi, gastral segments 1-2, 5-6, reddish, tergite 2 with two small round yellow spots. Mesoscutum with not very dense, dull greyish tomentum. Angola B.jordani sp. n. (p. 83) 47(38) Mesoscutum with dense silvery tomentum 48 Mesoscutum with less dense, fine tomentum which is usually quite inconspicuous .... 51 48(47) Mesoscutum and scutellum with deep, rather large punctures. Propodeum with strong striae. Mesoscutum with very dense silvery tomentum. Whole frons, upper part of gena, antennae, mesosoma, legs, gaster behind the stalk of tergite 2, black. Wings blackish with slight purple reflection, length 16-5 mm. Hind femora with dense but rather short hairs beneath. Uganda B. punctilio sp. n. (p. 83) REVISION OF BELONOGASTER 39 Mesoscutum and scutellum with much smaller less deep punctures. Propodeum less strongly striate 49 49(48) Wings blackish or dark grey, length 17-0-22-5 mm. Stalk of second gastral tergite 2-5 times as long as broad. Dark ferruginous, mesoscutum and gastral segments 3-5 blackish, 2 with large whitish yellow triangular spots. Hind femur with a few pale bristles beneath. Sudan (Kordofan, Dhafur), Congo (northern) B. fuscipennis du Buysson (p. 86) Wings red- or yellow-brown, tips darker, length 16-0-22-5 mm. Stalk of second gastral tergite 2-5-3-5 times as long as broad 50 50(49) Gaster often with 3 pairs of yellow spots in West African specimens but sometimes with 1 pair only on tergite 2, or, in South African specimens often with none. Legs often reddish with dark tarsi. Wings usually rather pale with tips darkened, length 16-0-22-5 mm. Antennal segment 3 nearly = 4 + 5 + half 6. Stalk of second gastral tergite 2-5-3-5 times as long as broad. Mid and hind femora with practically no bristles beneath. Chad, West, Central and South Africa B. grisea (F.) (p. 87) Gaster usually with no yellow spots, rarely a pair on tergite 2. Legs black or ferruginous with mid and hind tibiae and tarsi black. Wings red-brown, tips infuscate, length 18-0-20-0 mm. Antennal segment 3 nearly = 4 + 5 + 6. Stalk of second gastral tergite 2-5 times as long as broad. Hind femur with a few short bristles beneath. Head and mesosoma ferruginous, mesoscutum black with tomentum hiding the cuticle, gaster with very dense silvery tomentum posteriorly, cuticle ferruginous with tergites 3-6 more or less blackened. Kenya, Tanzania B. neavei sp. n. (p. 89) 51(47) Hind femora with numerous short black bristles beneath. Stalk of second gastral tergite 1-5-2-5 times as long as broad. Fifth segment of fore tarsus long. Mesoscutum blackish with at most indistinct punctures. Wings with dark tips, length not more than 19-5 mm. Gaster with no yellow markings 52 Hind femora without black bristles beneath or (B. brevitarsus) stalk of second gastral tergite 2-5-3-5 times as long as broad and fifth segment of fore tarsus short 53 52(51) Stalk of second gastral tergite 2-0-2-5 (rarely 3-5) times as long as broad. Gena 1-5 times as wide as eye in profile. Mesosoma granulate, not punctured. Legs dark ferruginous, tarsi black. Wings dark grey, length 16-5-19-0 mm. Sierra Leone, Gabon, Congo B. leonina sp. n. (p. 90) Stalk of second gastral tergite 1-5 times as long as broad. Gena 1-5 times as wide as eye in profile. Mesosoma with a few punctures on the mesoscutum and many fine ones on the mesopleuron. Legs ferruginous, tibiae and tarsi black. Wings dark reddish brown, length 19-5 mm. Congo B. barbata sp. n. (p. 92) 53(51) Stalk of second gastral tergite not more than 2, usually 1-5 times as long as broad. Hind femur with a few short white bristles beneath. Gastral tergite 2 usually with yellow spots. Mesoscutum with close brownish tomentum 54 Stalk of second gastral tergite twice as long as broad or longer 55 54(53) Stalk of second gastral tergite usually yellowish, twice as long as broad. Hind femora ferruginous. Gaster with two large comma-shaped yellow spots on tergite 2. Thorax and gastral tergites 3^4 usually more or less blackened. Wings rather pale. Zambia, Zim- babwe, Malawi, Tanzania, Mozambique, South Africa .... B. brachystoma Kohl (p. 68) Stalk of second gastral tergite usually ferruginous, rarely more than 1-5 times as long as broad. Hind femora usually blackened at base. Gaster with or without comma-shaped yellow spots on second tergite. Thorax and gastral tergites 3-4 more often blackened. Wings usually rather dark. Clypeus strongly reticulate above. Apex of gastral petiole usually with two pale dots. Kenya, Zambia, Zimbabwe, Malawi, Tanzania, Botswana, South Africa B. freyi du Buysson (p. 69) 55(53) Stalk of second gastral tergite 3-0-3-5 times as long as broad. Fifth segment of fore tarsus short. Mid and hind legs blackish, hind femora beneath with some black bristles. Gaster normally without yellow spots, often with some black hairs protruding through the tomentum (cf. couplet 12). Kenya, Uganda B. brevitarsus sp. n. (p. 92) Stalk of second gastral tergite not longer than 2-5 times as long as broad, or, in B. longitarsus sometimes as long as 3-5 times but then the fifth segment of the fore tarsus is long ... 56 56(55) Mesoscutum dull coal-black with very distinct punctures, especially in front. Mesopleuron closely and coarsely punctured. 40 O. W. RICHARDS Scutellum and metanotum red, contrasting with the mesoscutum and propodeum. Stalk of second gastral tergite 2 as long as broad. Legs ferruginous, each femur with a line of white pile beneath. Gaster black, segments 1-2 red, tergite 2 with two elongate, trans- verse yellow spots, second sternite with two smaller yellow spots. Wings dark purplish brown, length 16-0- 18-0 mm. Socotra B. saussurei Kirby (p. 93) Thorax ferruginous or, if black, much less strongly punctured 57 57(56) Thorax black, scutellum and metanotum sometimes red. Stalk of second gastral tergite 2- 5 times as long as broad 58 Thorax ferruginous 59 58(57) Legs and thorax entirely black. Wings purplish black, length 20-0 mm. Femora with a line of white pile beneath. Ethiopia B. abyssinica du Buysson (p. 96) Legs ferruginous, scutellum and metanotum red. Wings yellow-brown, tips very dark fus- cous, length 17-0-20-0 mm. Femora beneath only with white tomentum. Second gastral tergite sometimes with large triangular yellow spots but the yellow some- times more or less completely invaded by ferruginous. Somalia B. adenensis somaliensis subsp. n. (p. 100) 59(57) Wings dark purplish brown, length 19-0-21-0 mm. Femora with lines of not very dense white pile beneath. Thorax with numerous rather fine punctures. Propodeum striate behind. Gastral ter- gite 2 with transverse yellow spots, sternite 2 with spots or a band; gastral tergites 3-4 largely blackish. Oman (Dhofar) B. guichardi sp. n. (p. 94) Wings much paler, at least on proximal three-quarters. Femora without lines of white pile beneath 60 60(59) Gena 0-6-0-7 times as wide as eye in profile. Prementum and base of stipes beneath bare. Stalk of second gastral tergite 2-5-3-5 times as long as broad. Ferruginous, gastral tergite 2 with or without two round yellow spots. Uganda, Zambia, Angola, Mozambique, Tanzania, South Africa B. longitarsus sp. n. (p. 97) Gena 0-8-1-0 times as broad as eye in profile. Prementum and base of stipes with several long black bristles beneath. Stalk of second gastral tergite 2-0-2-5 times as long as broad. Second gastral tergite, if with yellow spots, with narrow transverse ones 61 61(60) Gastral tergites 3-4 black. Stalk of second gastral tergite 2-5 times as long as broad. Wings reddish brown with fuscous tips, length 17-0-20-0 mm. Gaster with no yellow spots. Southern Yemen (including Aden) . . . B. adenensis adenensis Giordani Soika (p. 99) Gastral tergites 3-4 ferruginous. Stalk of second gastral rather shorter. Tips of wings less darkened; wing-length 18-0-21-5 mm. (Two species often not cer- tainly distinguishable in $ except by the locality.) 62 62(61) Gastral tergite 2 with two narrow, very transverse spots at apex; sternite 2 also with two yellow spots or a band. Mesoscutum more strongly punctured. Southern Yemen, Yemen, SE. Saudi Arabia B. arabica Giordani Soika (p. 96) Gastral tergite 2 sometimes without yellow spots. Mesoscutum usually less strongly punc- tured. W. Saudi Arabia (around Jiddah) B.fitiformis (de Saussure) (p. 100) In B. adenensis, B. arabica and B. filiformis, the clypeus and malar space are sometimes yellowish. A female of B. arabica from Southern Yemen: W. Aden, Jebel Jihaf, 7,100 ft [2330 m], has the whole face yellow. Males No males have been seen of B. acaulis sp. n., B. atratus von Schulthess, B. aurata sp. n., B. barbata sp. n., B. ferruginea sp. n., B. flava sp. n., B. fuscipennis du Buysson, B. indica (de Saussure), B. jordani sp. n., B. kelnerpillautae sp. n., B. kohli Schulz, B. multipunctata sp. n., B. nigricans sp. n., B. punctilio sp. n., B. turbulenta Kohl, B. turgida Kohl. 1 Whole body including propodeum, smooth and shining. Small species, wing-length 12-0 mm. Clypeus very obtusely angled below. Antennal segments 4 and 5 four to five times as long as broad, segments 10-11 with a prominence at each end beneath, segment 12 about five times as long as broad. Nigeria (eastern province) B. nitida sp. n. (p. 60) Body dull with reticulation and tomentum. Species larger 2 2(1) Propodeum smooth and rounded, very finely granulate without striae, punctures, hairs or tomentum. REVISION OF BELONOG ASTER 41 Mesosoma without punctures and little tomentum. Stalk of second gastral tergite 1-5 times as long as broad. Gastral tergite 2 without yellow marks, posterior segments with short black bristles protruding. Wing-length 14-0 mm. Uganda, Liberia, Congo B. levior sp. n. (p. 58) Propodeum not smooth, without sculpture or vestiture 3 3(2) Clypeus below centrally only slightly protruding though obtusely angled, hardly protruding beyond the lateral lobes. Antennal segment 12 more or less blackened, flattened and shining 4 Clypeus below either little projecting but round or strongly pointed and projecting beyond the lateral lobes. Antennal segment 12 rarely blackened, flattened and shining .... 7 4(3) Clypeus below centrally with a slightly protruding blunt point but little bent down. Anten- nal segment 12 less flattened, narrower, usually blackened. Gena not quite as wide as eye. Hind trochanter and femur beneath with dense short white, and some longer blackish hairs. Very little yellow on clypeus, no yellow spots on gaster (B. brunnea Ritsema) ... 22 Clypeus with central process rounded. Antennal segment 12 strongly flattened, very broad, nearly always blackened beneath. Gena as wide or distinctly wider than eye. Hind tro- chanter and femur with longer and denser black or white hairs. Clypeus with yellow side-stripes, tergite 2 with yellow spots 5 5(4) Clypeus with very short, distinctly bent down central process. Gena much wider than eye. Hind femur and tibia with long hairs beneath. Gastral tergite 2 with narrow transverse yellow apical spots (B. clypeata Kohl) 6 Clypeus with somewhat longer, not bent down, rounded central process. Gena about as wide as eye or a little wider. Hairs of hind femur and especially of hind tibia fewer and shorter. Gastral tergite 2 with two large comma-shaped yellow spots. Zimbabwe, Zambia, Malawi, Tanzania, Mozambique, South Africa B. brachystoma Kohl (p. 68) 6(5) Colour paler ferruginous ; knees, especially the mid pair, yellowish. Sides of clypeus with little yellow. Hairs of femora and tibiae mostly white. Congo, Zambia, Uganda, Zim- babwe, Malawi, South Africa B. clypeata clypeata Kohl (p. 67) Colour darker, head beneath and frons black; knees not yellow. Sides of clypeus pale yellow. Long hairs of hind legs black. Zaire, Angola, Uganda, Zambia B. clypeata fuscata subsp. n. (p. 67) 7(3) Large (wing-length 22-0-25-0 mm), mainly black species, wings dark or black. Clypeus angled below but tip more or less rounded. Antennal segment 12 somewhat flattened, widened to apex, hardly curved 8 Smaller and not black or, if approaching this size and black, either the clypeus is widely rounded or pointed below, or antennal segment 12 is cylindrical and curved 11 8(7) Femora without pile or hairs beneath. Gaster without bristles among the tomentum 9 Femora beneath with pile, tomentum or hairs beneath. Mesoscutum more or less strongly punctured 10 9(8) Mesoscutum punctured, without hairs. Propodeum striate. Flagellar segments longer, seg- ment 8 2-1 times as long as broad. Kenya, Tanzania, Uganda, Zaire, Sierra Leone (?), Congo, Ivory Coast, Gabon, Chad B. saeva (de Saussure) (p. 61) Mesoscutum hardly punctured with very short hairs. Propodeum not striate. Flagellar segments shorter, segment 8 not quite twice as long as broad. Kenya, Malawi, Tanzania B. pileata sp. n. (p. 62) 10(8) All femora with a line of dense white tomentum beneath, hind femur beneath without hairs. Mesoscutum strongly punctate, without hairs. Propodeum striate. Gaster with black bristles amongst the tomentum. Congo, Zaire, Zambia, Uganda, Kenya, Zimbabwe, Angola, Malawi, Mozambique B. vasseae du Buysson (p. 56) Fore and mid femora with a line of dense white pile beneath, hind femora with rather less close white pile and many long black hairs. Mesoscutum very strongly and densely punctured and with long black hairs. Propodeum clathrate above, striate below. Gaster with very fine black bristles amongst the tomentum. Congo, Tanzania B. hirsuta sp. n. (p. 55) 11(7) Clypeus widely rounded below, not or scarcely pointed, not much projecting beyond the lateral lobes Clypeus below always with a pointed tip and usually produced well beyond the lateral lobes 24 1 2( 1 1 ) Fore and mid tarsi shortened and widened, segment 4 of mid tarsus much broader than long. Gastral tergite 2 with two yellow spots, stalk at least 3 times as long as broad. Gastral 42 O. W. RICHARDS petiole very long and slender. Thorax hairy. Antennal segment 1 1 with a strong protuber- ance beneath 13 Fore and mid tarsi longer and narrower (as usual or almost so), segment 4 of mid tarsus clearly longer than broad 14 13(12) Antennae with segments 1-8 darkened above. Mid tarsus moderately widened, segment 5 at least twice as long as wide. Legs not marked with yellow, mid and hind legs largely dark. Hind femur with dense white pile and scattered longer black bristles beneath. Fore tarsus very wide. Mid tibia without long hairs beneath. Kenya, Uganda B. brevitarsus sp. n. (p. 92) Antennae with segments not darkened above. Mid tarsus with segment 5 clearly less than twice as long as broad. Mid and hind legs largely ferruginous, fore and mid tarsi and mid femur anteriorly, mid tibia beneath, yellow. Hind femur beneath, especially towards the base, with long white or sometimes black hairs. Hind tibia with sparse long hairs beneath. Africa south of the Sahara B. filiventris (de Saussure) (p. 75) 14 Antennal segment 12 flattened and curved, usually blackened. Clypeus with ventral margin bisinuate. Stalk of second gastral tergite 1-5 times as long as broad. Antennal segments 1-7 more or less blackened above, segment 3 longer than 4 + 5. Clypeus with a slightly produced central point below. Hind femur with black and white bristles beneath. Gaster with no yellow spots. West Africa (cf. couplet 23) B. brunnea Ritsema (p. 64) Antennal segment 12 less flattened and curved. Clypeus smoothly curved from side to side. Stalk of second gastral tergite longer 15 15(14) Stalk of second gastral tergite 4-0-4- 5 times as long as broad 16 Stalk of second gastral tergite 1-5-2-5 times as long as broad 18 16(14) Gaster with no black hairs amongst the tomentum. Propodeum with strong striae and long outstanding silvery hairs. Cameroun 17 Gaster with fine black hairs amongst the silvery tomentum. Propodeum dull, granulate with mostly recumbent silvery grey hairs. Antennae dark above, 8-11 pale, 9-11 with flat projections beneath, 12 somewhat flattened, distal three-quarters shining black, tip narrowly rounded. Gastral petiole long and narrow. Wings brown, segments 2-3 with large yellow spots. Liberia B. libera sp. n. (p. 86) 17(16) Mesoscutum, humeri and most of mesopleuron blackish with strong quite close punctures. Antennae black above, segment 12 slightly flattened, straight below, curved above, seg- ments 10-11 long cylindrical, without prominences beneath. Gastral petiole very long and narrow. Wings light grey, length 19-5 mm. Gastral tergite 2 with no yellow spots. Cameroun B. punctata sp. n. (p. 84) Thorax ferruginous with weak, indistinct punctures. Antennae ferruginous, segment 12 shorter, more curved below, segments 10-1 1 short with a strong projecting keel below. Gastral petiole rather short, apical part distinctly widened. Wings hyaline, veins ferrugi- nous, length 17-0 mm. Gastral tergite 2 with two suboval almost contiguous spots. Cameroun B. rothkirchi von Schulthess (p. 85) 18(15) Large species, wing-length 19-0-28-0 mm. Gaster with short black bristles protruding amongst the tomentum. Clypeus with wide yellow side-stripes. Tips of wings hardly darkened. Frons ferruginous 19 Smaller species, wing-length 13-0-19-0 mm. Gaster without black bristles amongst the tomentum 20 19(18) Larger species, wing-length 22-0-28-0 mm. Antenna blackened above except segments 10-12, 12 sometimes more or less blackened but not strongly contrasting with adjoining segments, somewhat flattened, straight beneath, curved above, widening to apex. Tarsi black. Clypeus below bisinuate, central lobe regularly curved, a little projecting. West Africa, Zaire and Kenya to Tanzania, Malawi, Mozambique, South Africa . . . B. dubia Kohl (p. 53) Wing-length 22-0 mm. Antennae black above, segments 1-2 and extreme base of 3 yellow beneath, 3-9 ferruginous beneath, 10-11 light ferruginous, 12 blackish, not shining, a little flattened and curved, tip rounded, 10-1 1 cylindrical, not swollen beneath. Tarsi black, last segment of front pair reddish. Uganda B. ugandae sp. n. (p. 54) 20(18) Antennal segment 12 ferruginous, hardly shining with a few hairs, flat, straight below, curved above, tip rounded-truncate. REVISION OF BELONOGASTER 43 Antennal segment 1 1 cylindrical, 10 with a slight hump below. Mesoscutum with short, humeri with longer hairs. Face except for a rather narrow brown central stripe, light yellow, central lobe below regularly rounded. Hind femur beneath with tomentum and a few short bristles. Kenya, Uganda B. pusilloides sp. n. (p. 80) Antennal segment 12 shining, nearly always black, at least above, usually less flattened, more curved, tip rounded 21 21(20) Mesoscutum and humeri without outstanding hairs. Antennal segments 10-11 pale ferrugi- nous without prominences beneath. Clypeus with wide lateral white side-stripes, central lobe below just rounded, distinctly projecting. Hind femur beneath with long black and sometimes also some white hairs. Sierra Leone, Gabon, Congo B. leonina sp. n. (p. 90) Mesoscutum and humeri with distinct outstanding hairs or else antennal segments 10-11 with distinct prominences beneath 22 22(21) Frons brown. Antennal segment 12 shining black above, ferruginous below, much flattened and curved, 10-11 with considerable protuberances beneath. Clypeus almost regularly curved below, lateral white stripes quite wide. Hind femur beneath with white tomentum and a few short hairs. Guinea-Bissau, Sierra Leone, Ivory Coast, Ghana, Nigeria B. macilenta (F.) (p. 81) Frons black. Antennal segment 12 usually entirely shining black, rather less flattened, segment 1 1 cylindrical, 10 considerably thicker with a raised line beneath. All tarsi with fifth segment pale. All femora with long, rather dense hairs beneath (cf. couplet 14), (B. brunnea Ritsema) 23 23(22) Posterior part of gaster with less dense silvery tomentum. Wings distinctly reddish brown. Antennal segment 12 a little narrower. Kenya, Uganda B. brunnea nigriclava subsp. n. (p. 66) Posterior part of gaster with very dense silvery tomentum. Wings greyer. Antennal segment 12 a little thicker. Sierra Leone, Liberia, Ghana, Fernando Po, Congo B. brunnea brunnea Ritsema (p. 64) 24(1 1) Thorax and gaster with dense, brassy tomentum, hiding the cuticle. Antennae black, segment 12 much flattened, long oval, straighter below, more curved above, shorter than 11, dull; segments 10-11 with only raised line beneath. Clypeus obtusely pointed beneath, yellow with a narrow black central stripe. Coxae 1-2 yellow beneath, femora 1-2 with an anterior yellow spot at base. Wings dark brown, length 21-0 mm. Uganda, Burundi, Zaire B. leonhardii R. du Buysson (p. 87) Thorax and gaster without dense, brassy tomentum 25 25(24) Antennal segments 4 and 5 each clearly less than twice as long as broad. Antennae ferruginous, segment 12 curved, hardly flattened, longer than 5. Clypeus ven- trally, though pointed, not much protruding. Mesoscutum, scutellum, metanotum and propodeum with short, stout, curved bristles. Tibiae and tarsi ferruginous. Stalk of second gastral tergite not more than 1-5 times as long as broad. South Africa, Lesotho, Malawi, Zimbabwe, Tanzania B. petiolata (Degeer) (p. 71) Antennal segments 4 and 5 each more than twice as long as broad 26 26(25) Thorax and propodeum dorsally with dense, relatively long, black hairs. Antennae except segments 1-3 dorsally, ferruginous; segment 12 a little flattened and curved, as long as 9; 9-11 elongate, not much prominent beneath. Legs black, femora and tibiae 1-2 or 1-3 with pale yellow stripes beneath. Gaster with more or less distinct protruding black bristles, stalk of second gastral sternite 2-0-2-5 times as long as broad. Ethiopia B. meneliki Gribodo (p. 52) Thorax and propodeum without such dense long black hairs 27(26) Stalk of second gastral tergite not more than 1 • 5 times as long as broad. Antennae usually entirely ferruginous Stalk of second gastral tergite 2-3 times as long as broad 31 28(27) Gaster posteriorly with short black bristles protruding from the tomentum. Clypeus usually not or little yellow-marked; body except the head usually black or very dark ferruginous. Wings light to dark brown, tips a little darker, length about 19-0 mm. Antennal segment 12 moderately flattened and curved, about as long as 9. Kenya, Uganda, Zambia, Angola, Zaire B. pennata sp. n. (p. 51) Gaster without bristles in the tomentum or in B. lateritia only with very short ones. Clypeus with whitish or yellow side-stripes. At least the legs ferruginous. Wings yellowish brown, with dark tips (except in B. tarsata), length 16- 5- 19-0 mm 29 44 O. W. RICHARDS 29(28) Tarsi shortened, mid and hind pairs shining, without any hairs or bristles. Two small spots on gastral tergite 2, yellow. Wings light fuscous, length 17-5 mm. Stalk of second gastral tergite as long as broad. Thorax and propodeum black, former not punctured. Tanzania B. tarsata Kohl (p. 73) Tarsi not shortened, mid and hind pairs not so shining with numerous small bristles . . 30 30(29) Antennal segment 12 slightly flattened, distinctly curved, as long as segment 9; 10-11 with gently raised elongate prominences. Second gastral tergite sometimes with narrow yellow spots. Mesoscutum, mesopleuron and propodeum, gastral tergites 3-7 more or less black- ened; segments 1-2 ferruginous. Kenya, Tanzania, Zambia, Angola, South Africa B. freyi R. du Buysson (p. 69) Antennal segment 12 cylindrical, strongly curved, at least as long as 5; 10-11 not all raised beneath. Gastral tergite 2 with a large yellow spot. Small area on the frons, gastral tergites 4-5 more or less blackened. Kenya, Zambia, Zimbabwe, Angola, Malawi, Mozambique, South Africa . . . B. lateritia Gerstaecker (p. 72) 31(27) Gaster posteriorly with short black bristles protruding amongst the tomentum. Antennae more or less blackened, segment 12 more or less flattened and curved, stalk of second gastral tergite 2-0-2-5 times as long as broad. Gaster rarely with two yellow spots 32 Gaster posteriorly without black bristles amongst the tomentum, except sometimes at the extreme sides 34 32(31) Antennal segment 12 wider, more strongly flattened, moderately curved, especially above, as wide as segment 11 but a little longer than it; segments 9-11 with strong rounded protuberances beneath. Thorax and propodeum ratHr strongly punctured with relatively long black hairs. Clypeus sometimes not white at sides. Wings a little darkened. Kenya, Zambia, Tanzania, Malawi, South Africa (Natal, Orange Free State), Niger B. somereni sp. n. (p. 50) Antennal segment 12 narrower, less flattened, more curved and rather longer, as long as 9; 9-11 beneath with elongate, more or less raised protuberances beneath. Thorax without hairs. Clypeus white at sides (B. juncea (F.)) 33 33(32) Wings black with metallic reflections. Arabia, Algeria (Tripoli) (? strays), West Africa, Zaire, Uganda, Kenya, Malawi, Sudan B. juncea juncea (F.) (p. 48) Wings brownish with tips darker. Zaire, Uganda, Kenya, Malawi, Sudan B. juncea coloniatis Kohl (p. 48) 34(31) Wings dark brown, tips hardly darker, veins reddish, length 17-0-18-0 mm. Antennal segment 12 flattened, dorsal side curved, ventral side straight. Antennae and legs ferruginous 35 Wings less dark and their colour usually less uniform 37 35(34) Whole mesosoma ferruginous. Mesoscutum and pleuron less punctured. Antennal segment 12 shorter, 9-11 cylindrical with stronger ventral prominences whose surfaces are curved. Apex of gastral tergite 2, whole of 3, base of 4, blackish ; wide transverse spots on tergite 2, apical band of sternite 2, light yellow. Oman (Dhofar) B. guichardi sp. n. (p. 94) Mesosoma, except partly ventrally, black. Mesoscutum and pleuron more punctured. An- tennal segment 12 longer, 9-11 little projecting below, with flattened elongate promi- nences. Gastral tergites 3-4 black 36 36(35) Antennal segment 12 broader and flatter. Antennal scape not yellow beneath. Gaster with- out yellow markings. Mesopleuron less punctured. Ethiopia B. abyssinica du Buysson (p. 96) Antennal segment 12 narrower and less flattened. Antennal scape yellow beneath. Gastral segment 2 light ferruginous with apex of tergite and spots on sternite yellow. Socotra B. saussurei Kirby (p. 93) 37(34) Fore and mid tarsi shortened, segments 4 and 5 clearly shorter than on the hind pair. Antennal segment 12 long, curved, cylindrical, a little longer than 11. Antennal segments 10-1 1 with quite strong protuberances beneath. Ferruginous, meso- scutum and gastral segments 3-7 a little darkened. Clypeus with wide lateral white stripes ; gastral tergites 2-4 and sternite 2 each with a pair of yellow spots. Fore and mid femora and mid tibia with yellow stripes beneath. Wings hyaline, tips dark, length 15-5 mm. Kenya, Uganda, Zambia, Tanzania (including Zanzibar), Malawi, Mozambique B. facialis du Buysson (p. 80) Fore and mid tarsi not shortened. Antennal segment 12 shorter, less curved, less cylindrical 38 REVISION OF BELONOGASTER 45 38(37) Stalk of second gastral tergite 4-5 times as long as broad. Wings hyaline with ferruginous venation, length 17-0 mm. Largely ferruginous, femora somewhat darkened, gastral segments 2-7 blackish, 2 with two large contiguous yellow spots. Antennal segment 12 moderately flattened, a little curved above and below, segments 10-11 short with strongly protruding keels below. Cameroun B. rothkirchi von Schulthess (p. 85) Stalk of second gastral tergite 2-3 times as long as broad. Wings, except in the large B. brunnescens, with more or less distinct dark tips 39 39(38) Large species, length 20-0-22-0 mm, uniformly light brown. Uniformly ferruginous, at most the mesoscutum and gastral tergites 3-7 a little darkened. Stalk of second gastral tergite 2 as long as broad. Antennal segments 9-11 with slight convexities beneath. Ethiopia and Africa south of the Sahara. . . B. brunnescens sp. n. (p. 70) Smaller species or not uniformly ferruginous. Wings usually with distinct dark tips ... 40 40(39) Mesoscutum with dense silvery tomentum more or less hiding the cuticle. Gena clearly narrower than eye in profile 41 Mesoscutum with much finer and less dense tomentum, leaving the cuticle, which is very little punctured, exposed 43 41(40) Mesosternum with long black hairs in front of mid coxa. Mesoscutum and propodeum with dense, moderately long hairs. Antennal segment 12 as in B. grisea but rather shorter, segment 9 with a strong angular protuberance beneath. Femora 1 and 2 usually with yellow stripes beneath. Kenya, Tanzania B. neavei sp. n. (p. 89) Mesosternum without long black hairs in front of mid coxa 42 42(41) Antennal segment 12 cylindrical, as long as 8, slightly curved, segments 9-11 beneath with elongate, almost flat-topped elevations. Mesothorax (except scutellum), propodeum, gaster and mid and hind legs, practically black; mid and hind coxae beneath, strips on mid femur and tibia, large spots on gastral tergite 2, yellow. Zambia B. bimaculata sp. n. (p. 78) Antennal segment 12 somewhat flattened, as long as 8, more curved dorsally than ventrally; segments 8-9 with flat-topped elevation, 10-11 with angular protuberances, sometimes weak on 1 1. Mesothorax (except scutellum) usually more or less black, propodeum more ferruginous; base of gaster and femora ferruginous, only fore femur with a yellow stripe. Gastral tergite 2 with or without a pair of yellow spots. Chad, West, Central and South Africa B. grisea (F.) (p. 87) 43(40) Stalk of second gastral tergite usually twice as long as wide. Clypeus white, with a faint darker central stripe. Antennal segment 12 as long as 1 1, moderately flattened and curved; 10-11 with gently rounded protuberances beneath, prominence of 9 flatter. Thorax very feebly punctured. Ferruginous, sometimes gastral tergites 3-4 a little darker; sometimes narrow yellow spots at apex of gastral tergite 2. Wings reddish brown, tips a little darker. Yemen and Southern Yemen (Aden) B. arabica Giordani Soika (p. 96) Stalk of second gastral tergite 2-5-3-0 times as long as broad. Clypeal dark stripe more distinct 44 44(43) Ferruginous, gastral tergite 2 with two suffused yellow spots. Wings pale yellow-brown, length 15-0 mm. Antennal segment 12 moderately flattened, curved above, straight below, about as long as 7; 9-11 with strong but rounded protuberances beneath. Uganda, Zambia, Angola, South Africa, Tanzania B. longitarsus sp. n. (p. 97) At least gastral tergites 3-4 more or less darkened. Wings dark 45 45(44) Antennal segment 12 cylindrical, curved, very little flattened; segment 11 with an angular prominence. Antennal segment 9 with a flat prominence, 10 with an angular one. Ferruginous, gastral segments 3-7 usually blackish ; apparently tergite 2 sometimes with yellow spots. SW. Saudi Arabia B.fiUformis (de Saussure) (p. 100) Antennal segment 12 rather more flattened and less curved, 1 1 with a feeble prominence . . 46 46(45) Wings blackish. Much of gaster blackish, segment 2 without yellow spots. Head dorsally, mesosoma, blackish. Clypeus with wide lateral white stripes. Ethiopia B. abyssinica R. du Buysson (p. 96) Wings considerably paler. Gastral segments 3-7 black, segment 2 with narrow yellow spots (B. adenensis Giordani Soika) 47 47(46) Mesoscutum and propodeum sometimes darkened. Whole of clypeus except ventral margin 46 O. W. RICHARDS whitish yellow. Wings yellow-brown, tips light fuscous. Yemen and Southern Yemen (Aden) B. adenensis adenensis Giordani Soika (p. 99) Mesothorax (except pronotum, scutellum, metanotum and propodeum) black. Wings yellow-brown, tips dark fuscous. Somalia . . . . B. adenensis somaliensis subsp. n. (p. 100) Key to Malagasy species of Belonogaster 1 Stalk of second gastral tergite half as long as broad or a little longer. Gastral petiole swollen distally. Black; wings pale brown, length 18-0-20-0 mm. Female antenna with segments 4, 5 and 6, 1-1, 1-0 and 0-9 times as long as broad. Male mandibles and clypeus with dense outstanding white pubescence, clypeus below feebly produced; antennal segment 12 el- ongate, as long as 8, subcylindrical, quite strongly curved with numerous short hairs, 4-9 black, 10-12 light brown, 9-12 successively a little thinner B. brevipetiolata de Saussure (p. 101) Stalk of second gastral tergite at least clearly longer than broad. Gastral petiole not or only very gradually swollen distally 2 2(1) Very large, wing-length 25-0-29-0 mm. Disk of frons with outstanding black bristles. Genal margin to near or beyond the bottom of the eye. Practically no bristles beneath the femora. Tomentum very sparse, denser on posterior gaster but there very fine. Uniform brown, $ stalk of second gastral tergite 2-5, of <$ 1-7 times as long as broad. Male antennal segment 12 moderately flattened and curved, black, a little shining beneath B. guerini (de Saussure) (p. 102) Smaller, wing-length not more than 2 1-0 mm 3 3(2) Stalk of second gastral tergite 3 -5-4-0 times as long as broad 4 Stalk of second gastral tergite not more than 3 times as long as broad 6 4(3) Females (unknown in B. keiseri). Light brown, clypeus and legs beyond the coxae rather paler. Wings brownish hyaline, length 11-0-14-Omm. Disk of frons with rather fine outstanding brown bristles. Genal margin ending rather before the bottom of the eye. Tomentum very sparse, denser on posterior gaster but very fine. Femora with fine hairs beneath. Scutellum, metanotum and propodeum with numerous fairly long brown hairs. Valves of propodeum pale yellowish with a transparent, oval 'eye'-mark above it B. madecassa (de Saussure) (p. 103) Males 5 5(4) Yellowish ferruginous, head, legs and gaster beneath yellower. Wings hyaline, length 11 -Omm. Clypeus below gently rounded with moderately long and dense white hairs. Mesoscutum and femora beneath with no tomentum. Antennal segments 4-5 about 3 times as long as broad; 12 short (not quite as long as 8), flattened, a little widened to apex, tip narrowly black with a few short hairs; 8-11 beneath with protruding flat, table-like projections B. madecassa (de Saussure) (p. 103) Brownish ferruginous, most of mandibles, face below antennal sockets, legs except coxae, more or less dark brown. Malar space and gena, band across pronotal keel, lateral sclerites of scutellum, most of metanotum, valves and large posterior area on propodeum, mesopleural spot, pale yellow; fore coxa and femur, mid femur, tibiae and tarsi with creamy yellow marks. Wings light brownish hyaline, length 11 -5 mm. Clypeus projecting below in a rounded obtuse angle with dense outstanding hairs at sides. Mesoscutum with some long hairs, femora with fine pile rather than tomentum beneath. Antennal segments 4-5 about 3-5 times as long as broad, 12 narrow, considerably flattened, as long as 11 which is cylindrical, 8 a little more than 2-5 times as long as broad, 9-12 considerably longer and thinner, 6-10 with raised lines beneath B. keiseri sp. n. (p. 104) 6(3) Wing-length 17-0-2 1-0 mm. Stalk of second gastral tergite usually 3 times as long as broad, 2-5 times in B. apicalis $ 7 Wing-length 10-0- 16- 5 mm. Stalk of second gastral tergite not more than 2-5 times as long as broad. Mesoscutum granulate, not punctured 10 7(6) Bristles of mesosternum and fore coxa beneath, black. Antennal segment 3 clearly longer than 4 + 5. Wings yellowish hyaline, length 20-0 mm. Mid and hind femora (in good condition) with numerous hairs beneath. Ferrugi- nous; scape greenish black, coxae, femora, sometimes fore tibiae, black; gaster black, tergites 4-6 more or less ferruginous suffused. Mid and hind tibiae and sometimes fore REVISION OF BELONOGASTER 47 tibia, all tarsi, ochreous. Stalk of second gastral tergite blackish, slightly shorter than in B. prasina. Male not seen B. bicolor de Saussure (p. 104) Bristles of mesosternum and fore coxa pale or at most partly brown 8 8(7) Mid and hind femora with no hairs beneath. Wing-length 17-0-2 1-0 mm. Female antennal segment 3 a little longer than 4-1-5. Light ferruginous, frons, humeri, part of mesoscutum and sometimes scutellum, gastral petiole and anterior part of tergites 2-3, part of mid and sometimes hind femora, blue-green. Male with sides of clypeus, base of mandibles with dense outstanding white hairs. Clypeus roundly produced below. Antenna with segments 1-9 light brown, 10-12 black, 8- and 9 black marked beneath, segment 12 shining, curved, subcylindrical, 10-11 with a strongly raised ridge beneath projecting a little at apex. Fore and mid tarsus with seg- ments rather short and broad B. prasina de Saussure (p. 107) Mid and hind femora with quite conspicuous hairs as well as tomentum beneath. Wing- length 16-5- 18-5 mm. Males not seen 9 9(9) Antennal segment 3 distinctly longer than 4 + 5. Wings dark grey with tip largely light red-brown. Black, mandibles pitchy, ventral quarter of clypeus dark ferruginous. An- tennae ferruginous, segments 1-3 darkened above. Tarsi more or less reddened. Stalk of second gastral tergite more or less ferruginous . . . . . . B. apicatis de Saussure (p. 105) Antennal segment 3 a little longer than 4 + 5. Wings red-brown, tips if anything, paler. Ferruginous, frons, mesosoma, coxae and femora, gastral petiole and stalk of second gastral tergite somewhat darkened. Gaster sometimes vaguely darkened B. maromandia sp. n. (p. 108) 10(7) Females 11 Males 12 11(10) Hind tibia pale yellow with apex black. Mesosoma with surface granulate, not punctured, ferruginous, large part of humeri, mesoscutum and upper part of propodeum, black. Gaster black, tergites 2-6 with yellow apical bands spreading along the sides. Four hind coxae, all femora, four hind tarsi, black ; tibiae yellow. Wings very pale brownish hyaline, length 12-0- 13- 5 mm B. hildebrandti de Saussure (p. 108) Hind tibia of uniform colour, usually black but sometimes reddish. Thorax generally with a few distinct punctures, probably always some on pleuron. Colour usually light reddish brown but there is also a form which is largely blackish. Gaster usually not banded, but rarely a linear band on tergite 2. Wings hyaline, length 13-0 mm B. eumenoides de Saussure (p. 110) 12(10) Wings hyaline, length 12-0 mm. Anterior streaks on all femora, all tibiae, traces of bands at sides of gastral tergites 3-4, yellowish. Stalk of second gastral tergite about 2-5 times as long as broad. Antennae a little shorter, segments 8-12 shining beneath, 12 cylindrical, curved, as long as 8. Gastral petiole with distal part distinctly widened B. hildebrandti de Saussure (p. 108) Wings light brown, length 12-0 mm. Head light ferruginous brown, mandibles, malar space, narrow streak on gena, wide stripes from mid sinus to bottom of clypeus, whitish yellow. Spot above antennal sockets, antennal segments 1-2 beneath, basicostal plate, anterior streaks on femora 1-2, anterior streaks on all tibiae, narrow end and sides of gastral tergite 2, narrow end of sternite 2, broad streaks on coxae beneath, meso- and metaster- num, broadly, tip of pronotum below, creamy. Mid and hind tibiae blackish brown above. Stalk of second gastral tergite 2 as long as broad. Antennae longer, 6-10 with feeble raised lines beneath, 9-1 1 not shining beneath, segment 12 of similar shape but longer, as long as 9, traces of a division (into 12 and 13) rather before the middle. Gastral petiole with distal part widened B. eumenoides de Saussure (p. 110) Descriptions of African species Belonogaster juncea (F.) (Figs 1-5) Vespa juncea Fabricius, 1781 : 468. Raphigaster junceus (Fabricius) de Saussure, 1853: 14, pi. 2, fig. 2. Belonogaster junceus (Fabricius) Gerstaecker, 1862: 468. 48 O. W. RICHARDS This is a common species in tropical Africa and occurs in two forms, the nominotypical one with darker wings in more northern parts and a subspecies with lighter wings in the south. De Saussure includes B. juncea in his conspectus of the species of Madagascar but I know of no reliable record from that island. Belonogaster juncea juncea (F.) Vespa juncea Fabricius, 1781 : 468. Holotype 9, AFRICA AEQUINOCTIALI (Banks coll., BMNH) [examined]. IVespa guineensis Fabricius, 1793: 277. Holotype 9, GUINEA (Isert coll., Kiel) [not examined]. FEMALE. Head ferruginous; antennal segments 1-7 darkened above. Mesosoma ferruginous, dorsal part of humeri and mesoscutum darkened. Legs ferruginous, tibiae 2-3 a little darkened, tarsi black. Gaster dark ferruginous, tergites 2-6 gradually more blackened. A very few 9 in West Africa have two small yellow spots on gastral tergite 2. Wings black with purple reflections, length 17-0-23-5, mean (1569) 20-35 mm. Clypeus (Fig. 3) acute below, ventral quarter shining with scattered large punctures bearing short black bristles, dorsal part finely granulate, dull with more but smaller punctures and with very short black bristles and not very dense white tomentum. Frons with fairly close small punctures and outstanding black bristles, also rather sparse white tomentum. Gena not quite as wide as eye in profile. Antenna (Fig. 1) with segment 3 clearly longer than 4 + 5, 4 and 5 about 1-5 times as long as broad, 8 quadrate. Base of submentum and stipes with a few short black bristles. Mesoscutum and humeri finely granulate dull, with many small punctures and close silvery tomentum, mesopleuron similar; scutellum and metanotum with rather closer and stronger punctures, former with a strong impressed central line. Propodeum (Fig. 4) dull, very finely granulate, dorsally striate, sides punctured, tomentum close and silvery, bristles outstanding, short and black, posterior depression nearly half the dorsal length of propodeum, impressed line strong for two-thirds of length, anterior depression rather large, shallow. Last segment of the foretarsus long. Femora beneath with pale tomentum and not many short outstanding black or brown bristles. Gastral petiole relatively short and stout, a little widened at apex, spiracles little projecting, hairs sparse. Stalk (Fig. 5) of second gastral tergite 2-3 usually 2-5 times as long as broad. Gaster posteriorly with rather close pale tomentum through which numerous short black bristles protrude. MALE. Very like the female; dorsal streak on mandibles, sides of clypeus, inner orbits, anterior streak on scape, yellow. Two males with two small yellow spots on gastral tergite 2. Wing-length 15-5-22-0, mean (42