XIV. PATHOGENS OF GLOSSINA (TSETSE FLIES)a Richard A. Nolan Department of Biology Memorial University of Newfoundland St. John's, Newfoundland, Canada AIC SS7
a This study was supported by National Research Council of Canada grant A-6665.
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271
ABSTRACTS
Richard A. Nolan
Bacterial infections in Glossina morsitans Westwood fed through membranes. Bauer, B. (1974). (Abstr.) Proc. 3rd Int. Cong. Parasitol. (Munich), 2: 943.
Pseudomonas and Aeromonas were isolated from dead and dying flies with black abdomens. Transmission of the bacteria by saliva of artificially infected flies was demonstrated. Most died within 8 days. An account of the biology of The natural history of tsetse flies. Buxton, P. A. (1955). Lond. Sch. Hyg. Trop. Med. Mem., No. 10, 816 pp. the genus Glossina (Diptera). Also reference is Buxton reports a Leptomonas sp. infecting G. morsitans in Nigeria. made to an observation by Hoare in 1932 of G. palpalis infected by Hepatozoon pettiti. The tsetse had acquired the haemogregarine by feeding upon Crocodilus niloticus which It was assumed that transmission of was infected with Haemogregarina pettiti Thiroux. This was the haemogregarine occurred when a fly was crushed in the crocodile's mouth. the first report definitely establishing a Dipteran as an intermediate host for a
haemogregarine. Progress report on investigations into the bionomics of Carpenter, G. D. H. (1912). Rep. Sleeping Sickness Bur. Lond., 12: Glossina palpalis, July 27, 1910 to August 5, 1911. 79-111.
The fly was originally reported as being G. palpalis, but this error was corrected in Gut contents of 455 Carpenter (1913), and the identification changed to G. fuscipes. An unidentified ascomycete with hyaline ascospores, a member of flies were examined. the Fungi Imperfecti, and filaments of an unidentified alga were infrequently observed Also, a larval nematode was found coiled in the gut of a single in the gut contents. Bacilli were also observed and Carpenter felt that there was fly just after emergence. a marked incompatibility between bacilli and trypanosomes in the gut of the fly. Second report on the bionomics of Glossina fuscipes (palpalis) of Carpenter, G. D. H. (1913). Rep. Sleeping Sickness Bur. Lond., 14: 1-37. Uganda.
When large numbers of bacilli were present, the fore-gut wall seemed thin and friable, Also, 4 nematodes were and Carpenter felt it possible that the flies were "sickly". found in the coelomic cavity of separate flies of over a thousand flies dissected.
Sporogonie d'une hemogregarine chez une ts6tsd (Glossina Chatton, E. & Roubaud, E. (1913). Bull. Soc. Pathol. Exot., 6: 226-233. palpalis R. Desv.). The first report of a haemogregarine from Glossina. Doidge, E. M. (1950). 1-1094.
It was found in the coelomic cavity.
The South African fungi and lichens to the end of 1945.
Bothalia, 5:
Recognized as a valid fungal taxon, Fusarium semitectum Berk. and Rav. is considered to be extremely common in tropical and subtropical countries and to be a secondary invader of However, some diseases of animals are plant tissue and seldom found to be pathogenic. produced by toxin-producing species of Fusarium (fusariotoxicoses).
272
Dutton, J. E. et al. (1907). Concerning certain parasitic protozoa observed in Africa. Part II. Ann. Trop. Med. Parasitol., 1: 287-370.
Myxosporidium heibergi should be considered of uncertain affinities due to the incomplete nature of the study. Infestation of Glossina palpalis R.-D. 1830 (Diptera) by larval Foster, R. (1963). Mermithidae Braun 1883 (Nematoda) in West Africa, with some comments on the parasitization of man by the worms. Ann. Trop. Med. Parasitol., 57: 347-358.
Of 4001 G. palpalis collected in Liberia, 15 were infested with larval mermithid. The nematodes were found in the abdominal cavity lying loosely intertwined with the gut and This article is a good review of nematode records through Malpighian tubules. rou Thomson (1947). Adults of G. fusca Walker, G. pallicera Bigot and C Foster felt Newstead were also examined and found to be free of nematode infections. The 15 nematodes infecting that the tsetse became infected while resting on the ground. G. palpalis were found in the coelomic cavity. Foster, R. (1964). An unusual protozoal infection of tsetse flies (Glossina Weidemann 1830 spp.) in West Africa. J. Protozool. 11(1): 100-106.
The flagellate was similar to, but not synonymous with Trypanosoma grayi Novy (1906). This unidentified flagellate accounted for approximately 90% of the total flagellate infections. The infection was usually confined to the midgut with hind-gut infections being rare. In about 8.5% of the infections, the flagellates occurred in the coelomic cavity only; and in approximately 8.57, of the infections, they occurred in both the coelomic cavity and the midgut. This is the first report of a trypanosome developing in the coelomic cavity of tsetse; the mouthparts were never invaded. The production of "symbiont-free" Glossina morsitans and an Hill, P. et al. (1973). associated loss of female fertility. Trans. R. Soc. Trop. Med. Hyg., 67: 727-728.
This reference is included in the coverage of the literature because it has implications The microbial control envisaged here for the use of microbial control of tsetse. involves the elimination or reduction of a microorganism by chemical means in order to reduce the level of tsetse instead of the normal concept of introducing and/or increasing the level of a pathogenic microorganism.
Huebner, E. & Davey, K. G. (1974). 249 (5454): 260-261.
Bacteroids in the ovaries of a tsetse fly.
Nature,
Bacteroids were present in the cytoplasm of oocytes 'and nurse cells of the ovary and The authors suggest that transovarian occasionally in follicle cells and sheath cells. transmission of the symbiont may occur.
Virus-like particles in a strain of G. morsitans centralis, Machado Jenni, L. (1973). Trans. R. Soc. Trop. Med. Hyg., 67(2): 295.
(1970).
The VLP were found in cytoplasmic vesicles of salivary glands and were first seen in The VLP were composed of an electron-dense core flies four days after emergence. The approximately 240A across and a surrounding envelope measuring 500-550A in diameter. VLP were also associated with special rod-like structures. This study and the discovery of a VLP in G. fuscipes fuscipes fienni, L. & Steiger, R. (197427 point out the need for established, susceptible tsetse cell lines to allow viral replication in vitro in order that a proper evaluation of viruses from Glossina can be made.
273
Jenni, L. & Steiger, R. (1974). Viruslike particles in Glossina fuscipes fuscipes Newst. Acta Trop., 31: 177-180. 1910. The viruslike particles measure about 350-390A in diameter and form intranuclear pseudocrystalline arrays in midgut epithelial cells. Young flies were found to have disrupted midguts, and the occurrence of VLP aggregates in older flies could be regarded as a latent infection. There was a high mortality rate in young flies (within 10 days after emergence) and a low hatching rate.
Leiper, R. T. (1910). 147.
L-xhibition
of a series of Entozo.
Proc. Zool. Soc. Lond., 79:
A single immature female Mermis was observed infecting G. palpalis. three inches in length and from the coelomic cavity of the tsetse.
The specimen was
Mr Lloyd records a "New Nematode parasite of Glossina morsitans". Leiper, R. T. (1912). J. Lond. Sch. Hyg. Trop. Med., 2: 41-42. Four immature nematodes were found in the tsetse coelomic cavity. Two were in one fly. Leiper refers to a similar immature Mermis found previously by a Professor Minchin a few years prior to this report in Glossina palpalis in Uganda.
Lester, H. M. 0. (1934).
Report of the tsetse investigation.
Rep. Med. Health Serv.
Nigeria 1933, pp. 74-83. Flies infected with 2 unidentified fungi were found at the end of the wet season. Note on the occurrence of a herpetomonad in Glossina morsitans. Lloyd, L. (1924). Ent. Res., 15: 185-186.
Bull.
The flagellate was found in the proboscis and in the midgut. This organism was not encountered in 10 000 G. tachinoides examined by Lloyd's group. Wenyon, C. M. (1926, p. 434) speculated that this organism might be a Leishmania as human leishmaniasis occurs in Nigeria. Second report of the tsetse-fly investigation in the northern Lloyd, L. et al. (1924). Provinces of Nigeria. Bull-. Ent. Res., 15: 1-27. An unidentified reptilian haemogregarine was found in G. tachinoides, The incidence in the fly depended upon the reptilian host: crocodile, frogs, and colubrine snake, 42%; monitors and lizards (Varanus), 457/; and chameleon, 54%. No evidence of pathogenicity to G. tachinoides was found.
Ma, W.-C. & )enlinger, D, L. (1974). Secretory discharge and mieroflora of milk gland in tsetse flies. Nature, 247(5439): 301-303. The results in G. morsitans morsitans which indicated the presence of the posoibly gramnegative bacteria in the lumen of the milk gland were also confirmed by compArative observations of G. austeni Newstead and G. longipalpis pallidiees Atisten. This study indicates some ultrastructural differences between the bacteroids of the mycetome And the bacteria in the milk gland lumen.
274
Notes on some blood parasites collected in Nigeria. Macfie, J. W. S. (1914a). Med. Parasitol., 8: 439-465.
Ann. Trop.
One tsetse fly "out of a number examined" was infected by an unidentified spirochaete. Macfie felt that this spirochaete was distinct from Spirochaeta glossinae described Macfie kept the tsetse in captivity for 22 from G. palpalis by Novy & Knapp (1906). days before it was killed for examination, and the fly did not appear to be affected by the presence of the large number of active spirochaetes.
The occurrence of a spirochaete in the gut of Glossina tachinoides. Macfie, J. W. S. (1914b). The spirochaete was apparently non-pathogenic. Ann. Trop. Med. Parasitol., 8: 464-465.
The results of dissections of tsetse flies at Accra. Macfie, J. W. S. (1915). Lab. Lond., pp. 49-54, 98-99.
Rep. Accra
The Seventy-five flies were dissected and 3 possibly parasitic organisms were found. abdomen of one infected fly was swollen; and it was also hard and pale-coloured ReproDeath was felt to be due, without doubt, to a fungal infection. posteriorly. The hyphae formed duction of the fungus appeared to occur via arthrospore formation. a dense network in which the lower half of the gut and the testicles were firmly embedded. Spirochaetes were found in the gut of another fly and appeared to be morphologically identical to those found by Macfie in the gut of a specimen of G. tachinoides in Nigeria. Also, sporocysts were found in the abdominal cavities of 2 flies and were similar to those found by Chatton and Roubaud (1913) in the abdominal cavity of the same species. Some observations on the seasonal spread of Glossina pallidipes in Moggridge, J. Y. (1936). Italian Somaliland with notes on G. brevipalpis and G. austeni. Bull. Ent. Res., 27: 449466. An unidentified Phycomycete was found in G. brevipalpis. Moggridge noted that the fungus was similar to a pathogenic fungus known in certain areas of Tanzania and that the fungus was present on the abdomen of the flies.
Moloo, S. K. (1972).
Parasitol., 66:
Mermithid parasite of Glossina brevipalpis Newstead.
Ann. Trop. Med.
159.
Two mermithid larvae were found in the coelomic cavity of the one infected adult tsetse. The 5000 flies examined included G. fuscipes, G. brevipalpis and G. pallidipes. Moloo felt that the infection of Glossina was accidental.
The ecology of Glossina morsitans, Westw., and two possible methods Nash, T. A. M. (1933). for its destruction. Part I. Bull. Ent. Res., 24: 107-157. The article includes the statement that a Mr Scott did some work with a fungus and proved Infected larvae had black spots on that it was definitely pathogenic to G. morsitans. their abdomens. Control by parasites and predators of Glossina, pp. 521-532. Nash, T. A. M. (1970). In Mulligan, H. W. (ed.), The African Trypanosomiases, Allen & Unwin Ltd, London. A personal communication from W. H. Potts reports that infection by an unidentified Phycomycete neither prevented completion of development of tsetse larvae nor their deposition.
275 A nematode, tentatively Personal communication to R. F. Myers. Nickle, W. (1973). The identified as Hexamermis sp., was found in the coelomic cavity of G. morsitans. collection was made by S. N. Okiwelu. Three unidentified larval Personal communication to R. F. Myers. Nickle, W. (1974). The collection was made mermithid were found in the coelomic cavities of G. palpalis adults. by K. Riordan.
Investigations on the role of symbionts in tsetse flies (Glossina Nogge, C. (1974). morsitans). Proc. 3rd Int. Cong. Parasitol. (Munich) 2: 947. Attempts were made Symbiont-free flies were obtained by oral treatment with lysozyme. to compensate for the loss of the symbionts by feeding defibrinated blood supplemented by B vitamins. Other techniques such as the raising and lowering of the temperature, the feeding of blood with added sodium oxalate, 7-chlortetracycline, and lysozyme in different concentrations as well as the injection of lysozyme into the coelom resulted in The destruction of the symbionts as well as a decrease in the longevity of the flies. blood meals of these latter flies remained undigested. Novy, F. G. & Knapp, R. E. (1906). J. Infect. Dis., 3: 291-393.
Studies on Spirillum obermeieri and related organisms.
A bacterium, Borrelia glossinae, was described as Spirillum glossinae in 1906 from smears It is listed in Buchanan and Gibbons (Buchanan, R. E. & of flies' stomach contents. 8th Edition. Bergey's Manual of Determinative Bacteriology. 1974. Gibbons, N. E. Baltimore. p. 190) as a Species incertae sedis The Williams and Wilkins Company. because of inadequate description.
The occurrence of intracellular rickettsia-like Pinnock, D. E. & Hess, R. T. (1974). organisms in the tsetse flies, Glossina morsitans, G. fuscipes, G. brevipalpis and Acta Trop., 31: 70-79. G. pallidipes.
Rickettsia-like organisms were observed in electron microscopy preparations the midgut epithelium, including the mycetome, in cells associated with the in developing oocytes. The authors concluded that the presence of a lytic the organisms and that the disruption of cells at the ultrastructural level that the rickettsia-like organisms were parasitic.
of cells of fat body and zone around
suggested
Rapport sur les Travaux de la Mission Scientifique du Katanga Rodhain, J. et al. (1913). (Octobre 1910 a Septembre 1912). Hayez. Bruxelles. 258 p.
Single nematode larvae (possibly Mermis sp.) were found in 4 flies. examined was not given.
The total number
Microorganisms in spermathecae of wild Glossina pallidipes. Rogers, A. (1973). Soc. Trop. Med. Hyg., 67: 299.
Trans. R.
These microorganisms occurred in the lumen of the spermathecae in both inseminated and uninseminated females in wild populations and also occurred at high concentrations in the spermathecae of laboratory-reared females.
Roubaud, E. (1911). 122-132.
Etudes sur les Stomoxydes du Dahomey.
Bull. Soc. Pathol. Exot., 4,
Roubaud attempted to infect by means of direct contact with Entomophthora muscae: 8 G. palpalis, 4 G. tachinoides, and one G. longipalpis and by means of ingestion of the
276
spores: 10 Glossina palpalis. The Glossinas were not infected by either means. The sporangiospores used had been attached to potato leaves and had originated from an infected Stomoxys calcitrans L. However, E. Miller-Kogler (Pilzkrankheiten bei Insekten. Parey. Berlin. 1965, pp. 160-161) has previously pointed out the lack of controls in this study.
Roubaud, E. (1919). Les particularit6s de la nutrition et la vie symbiotique chez les mouches ts6tses. Ann. Inst. Pasteur, 33: 489-536.
The mycetome is located Cicadomyces sp. is a symbiote of G. palpalis and G. tachinoides. in the middle of the anterior segment of the mid-gut, where absorption but no digestion occurs (Wigglesworth, V. B. (1972). The Principles of Insect Physiology. Chapman & Hall. London, p. 523).
Roubaud, E. & Treillard, M. (1935). Un coccobacille pathogene pour les mouches Tsetses. C. R. Hebd. Seances Acad. Sci. (Paris) 201: 304-306. This microorganism, Bacterium mathisi, which initially showed promise as a biocontrol agent has neither been maintained in culture nor is it a recognized valid taxon. Roubaud & Treillard (1935) found that the bacterium was taken in via the proboscis during feeding on guinea pigs whose hair and skin had been coated with the bacterium. The flies are apparently killed by a septicemic infection within the coelom within 1-8 days. Several insects were shown to be susceptible when inoculated with bacterial suspensions; however, Culex pipiens, Stegomyia sp. and Aedes aegypti survived ingestion of the microbe.
Roubaud, E. & Treillard, M. (1936). Infection experimentale de Glossina palpalis par un coccobacille pathogane pour les muscides. Bull. Soc. Pathol. Exot., 29: 145-147. No naturally occurring infection by Bacterium mathisi was found in the several hundred Glossina palpalis shipped from Uganda in the pupal stage. The low number of flies used in the experiments makes interpretation difficult; however, it should be noted that a septicemia was not indicated in all cases of adult mortality. When G. morsitans from Tanzania were used in experiments, they died of a septicemia in 1 to 3 days after feeding on contaminated guinea pigs.
Swynnerton, C. F. M. (1936). The tsetse flies of East Africa. A first study of their ecology, with a view to their control. Trans. R. Entomol. Soc. Lond., 84: 1-580. An unidentified Phycomycete was found infecting G. palpalis. Material for the cultivation of the fungus was sent to Johannesburg; however, the study does not appear to have been continued. Swynnerton felt that the hope of success using this particular fungus was small because it seemed to be fatal only during periods of wet weather.
Thomson, W. E. F. (1947).
Nematodes in tsetse.
Ann. Trop. Med. Parasitol., 41:
164.
The nematodes were located in the coelomic cavity and had an average length of 79 mm. InterVago, C. & Meynadier, S. pp. 24, 27, In Strains of Entomophagous Microorganisms. WPRS Bulletin national Organization for Biological Control of Noxious Animals and Plants.
1973/3. Aspergillus flavipes and Penicillium janthinellum were isolated from G. fusca congolensis in 1965. Both fungi have been previously isolated from soil in Africa.
277 Vanderyst, P. H. (1923). 32-42.
La prophylaxie contre la trypanose humaine.
Rev. Med. Angola, 4:
A local fly "Nkulu nyansi" was very rapidly infected with Entomophthora muscae when placed in a tube with one of the infected M. domestica L. received from Europe; however, The experiment was later G. palpalis was not infected under the same conditions. supplemented by adding locally obtained M. domestica, and these were also not infected. One of the tsetses used did die during the course of the experiment but did not show any signs of a disease. The infection of the fly "Nkulu nyansill was felt to be enhanced by Other experiments with this local fly gave negative injury during insect procurement. results. This is the first attempt at assessing the virulence of a microbial pathogen derived from outside Africa. La transmission cyclique de races r6sistantes de van Hoof, L. & Henrard, C. (1934). Ann. Soc. Belge Med. Trop., 14: 109-144. Trypanosoma gambiense par Glossina palpalis.
Hyphae of an unidentified fungus occurred in the coelomic cavity, in the gut, in the musculature of the thorax and in the proventriculus of G. palpalis. These authors cite a reference (Lloyd & Johnson, 1922), in the text to a similar infection but do not include it in the bibliography. Etudes sur Recherches sur les champignons pathogenes pour les glossines. Vey, A. (1971). Rev. Elev. Med. Glossina fusca congolensis Newst. et Evans. en Republique Centrafricaine. Vet. Pays Trop. (N.S.) 24: 577-579. Two fungal species were isolated from tsetse pupae, Absidia repens and Penicillium Both are common soil inhabitants. Experimental infections of tsetse larvae lilacinum. are reported. Larvae were wounded prior to incubation with the spore suspensions.
An unidentified coccoid bacterium was Personal communication to R. Nolan. Vey, A. (1974). isolated from Glossina pupae at the Laboratory I.E.M.U.T. de Farcha (Chad). Aspergillus ochraceus was isolated from Glossina pupae into pure culture in the Laboratory I.E.M.U.T. de Farcha (Chad). A. ochraceus has also been found in material of Theobroma cacao from the Gold Coast (Dade, H. A. 1940. A revised list of Gold Coast Fungi and Plant Diseases. Bulletin of Miscellaneous Information No. 6, pp. 205Kew Bulletin. Royal Botanic Gardens.
247). Wallace, J. M. (1931). Parasitol., 25: 1-19.
Micro-organisms in the gut of Glossina palpalis.
Ann. Trop. Med.
A fairly thorough study which would indicate that G. palpalis does not readily pick up bacteria from contaminated surroundings, such as fly boxes; however, during feeding Wallace felt that some earlier workers experiments organisms can be introduced into the gut. had introduced bacteria which were surface contaminants into gut preparations during He also observed that in every instance where Trypanosoma grayi was present the dissection. This latter finding is contrary to those of Carpenter (1912). bacteroids were also present. Wallace's comments with regard to the general inability of G. palpalis to acquire bacteria within the gut by feeding through contaminated skin or hair because of a possible downward flow of saliva or because saliva is bactericidal is contrary to the findings of Roubaud & Treillard (1935) using G. morsitans.
a This study was supported by National Research Council of Canada grant A-6665.
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271
ABSTRACTS
Richard A. Nolan
Bacterial infections in Glossina morsitans Westwood fed through membranes. Bauer, B. (1974). (Abstr.) Proc. 3rd Int. Cong. Parasitol. (Munich), 2: 943.
Pseudomonas and Aeromonas were isolated from dead and dying flies with black abdomens. Transmission of the bacteria by saliva of artificially infected flies was demonstrated. Most died within 8 days. An account of the biology of The natural history of tsetse flies. Buxton, P. A. (1955). Lond. Sch. Hyg. Trop. Med. Mem., No. 10, 816 pp. the genus Glossina (Diptera). Also reference is Buxton reports a Leptomonas sp. infecting G. morsitans in Nigeria. made to an observation by Hoare in 1932 of G. palpalis infected by Hepatozoon pettiti. The tsetse had acquired the haemogregarine by feeding upon Crocodilus niloticus which It was assumed that transmission of was infected with Haemogregarina pettiti Thiroux. This was the haemogregarine occurred when a fly was crushed in the crocodile's mouth. the first report definitely establishing a Dipteran as an intermediate host for a
haemogregarine. Progress report on investigations into the bionomics of Carpenter, G. D. H. (1912). Rep. Sleeping Sickness Bur. Lond., 12: Glossina palpalis, July 27, 1910 to August 5, 1911. 79-111.
The fly was originally reported as being G. palpalis, but this error was corrected in Gut contents of 455 Carpenter (1913), and the identification changed to G. fuscipes. An unidentified ascomycete with hyaline ascospores, a member of flies were examined. the Fungi Imperfecti, and filaments of an unidentified alga were infrequently observed Also, a larval nematode was found coiled in the gut of a single in the gut contents. Bacilli were also observed and Carpenter felt that there was fly just after emergence. a marked incompatibility between bacilli and trypanosomes in the gut of the fly. Second report on the bionomics of Glossina fuscipes (palpalis) of Carpenter, G. D. H. (1913). Rep. Sleeping Sickness Bur. Lond., 14: 1-37. Uganda.
When large numbers of bacilli were present, the fore-gut wall seemed thin and friable, Also, 4 nematodes were and Carpenter felt it possible that the flies were "sickly". found in the coelomic cavity of separate flies of over a thousand flies dissected.
Sporogonie d'une hemogregarine chez une ts6tsd (Glossina Chatton, E. & Roubaud, E. (1913). Bull. Soc. Pathol. Exot., 6: 226-233. palpalis R. Desv.). The first report of a haemogregarine from Glossina. Doidge, E. M. (1950). 1-1094.
It was found in the coelomic cavity.
The South African fungi and lichens to the end of 1945.
Bothalia, 5:
Recognized as a valid fungal taxon, Fusarium semitectum Berk. and Rav. is considered to be extremely common in tropical and subtropical countries and to be a secondary invader of However, some diseases of animals are plant tissue and seldom found to be pathogenic. produced by toxin-producing species of Fusarium (fusariotoxicoses).
272
Dutton, J. E. et al. (1907). Concerning certain parasitic protozoa observed in Africa. Part II. Ann. Trop. Med. Parasitol., 1: 287-370.
Myxosporidium heibergi should be considered of uncertain affinities due to the incomplete nature of the study. Infestation of Glossina palpalis R.-D. 1830 (Diptera) by larval Foster, R. (1963). Mermithidae Braun 1883 (Nematoda) in West Africa, with some comments on the parasitization of man by the worms. Ann. Trop. Med. Parasitol., 57: 347-358.
Of 4001 G. palpalis collected in Liberia, 15 were infested with larval mermithid. The nematodes were found in the abdominal cavity lying loosely intertwined with the gut and This article is a good review of nematode records through Malpighian tubules. rou Thomson (1947). Adults of G. fusca Walker, G. pallicera Bigot and C Foster felt Newstead were also examined and found to be free of nematode infections. The 15 nematodes infecting that the tsetse became infected while resting on the ground. G. palpalis were found in the coelomic cavity. Foster, R. (1964). An unusual protozoal infection of tsetse flies (Glossina Weidemann 1830 spp.) in West Africa. J. Protozool. 11(1): 100-106.
The flagellate was similar to, but not synonymous with Trypanosoma grayi Novy (1906). This unidentified flagellate accounted for approximately 90% of the total flagellate infections. The infection was usually confined to the midgut with hind-gut infections being rare. In about 8.5% of the infections, the flagellates occurred in the coelomic cavity only; and in approximately 8.57, of the infections, they occurred in both the coelomic cavity and the midgut. This is the first report of a trypanosome developing in the coelomic cavity of tsetse; the mouthparts were never invaded. The production of "symbiont-free" Glossina morsitans and an Hill, P. et al. (1973). associated loss of female fertility. Trans. R. Soc. Trop. Med. Hyg., 67: 727-728.
This reference is included in the coverage of the literature because it has implications The microbial control envisaged here for the use of microbial control of tsetse. involves the elimination or reduction of a microorganism by chemical means in order to reduce the level of tsetse instead of the normal concept of introducing and/or increasing the level of a pathogenic microorganism.
Huebner, E. & Davey, K. G. (1974). 249 (5454): 260-261.
Bacteroids in the ovaries of a tsetse fly.
Nature,
Bacteroids were present in the cytoplasm of oocytes 'and nurse cells of the ovary and The authors suggest that transovarian occasionally in follicle cells and sheath cells. transmission of the symbiont may occur.
Virus-like particles in a strain of G. morsitans centralis, Machado Jenni, L. (1973). Trans. R. Soc. Trop. Med. Hyg., 67(2): 295.
(1970).
The VLP were found in cytoplasmic vesicles of salivary glands and were first seen in The VLP were composed of an electron-dense core flies four days after emergence. The approximately 240A across and a surrounding envelope measuring 500-550A in diameter. VLP were also associated with special rod-like structures. This study and the discovery of a VLP in G. fuscipes fuscipes fienni, L. & Steiger, R. (197427 point out the need for established, susceptible tsetse cell lines to allow viral replication in vitro in order that a proper evaluation of viruses from Glossina can be made.
273
Jenni, L. & Steiger, R. (1974). Viruslike particles in Glossina fuscipes fuscipes Newst. Acta Trop., 31: 177-180. 1910. The viruslike particles measure about 350-390A in diameter and form intranuclear pseudocrystalline arrays in midgut epithelial cells. Young flies were found to have disrupted midguts, and the occurrence of VLP aggregates in older flies could be regarded as a latent infection. There was a high mortality rate in young flies (within 10 days after emergence) and a low hatching rate.
Leiper, R. T. (1910). 147.
L-xhibition
of a series of Entozo.
Proc. Zool. Soc. Lond., 79:
A single immature female Mermis was observed infecting G. palpalis. three inches in length and from the coelomic cavity of the tsetse.
The specimen was
Mr Lloyd records a "New Nematode parasite of Glossina morsitans". Leiper, R. T. (1912). J. Lond. Sch. Hyg. Trop. Med., 2: 41-42. Four immature nematodes were found in the tsetse coelomic cavity. Two were in one fly. Leiper refers to a similar immature Mermis found previously by a Professor Minchin a few years prior to this report in Glossina palpalis in Uganda.
Lester, H. M. 0. (1934).
Report of the tsetse investigation.
Rep. Med. Health Serv.
Nigeria 1933, pp. 74-83. Flies infected with 2 unidentified fungi were found at the end of the wet season. Note on the occurrence of a herpetomonad in Glossina morsitans. Lloyd, L. (1924). Ent. Res., 15: 185-186.
Bull.
The flagellate was found in the proboscis and in the midgut. This organism was not encountered in 10 000 G. tachinoides examined by Lloyd's group. Wenyon, C. M. (1926, p. 434) speculated that this organism might be a Leishmania as human leishmaniasis occurs in Nigeria. Second report of the tsetse-fly investigation in the northern Lloyd, L. et al. (1924). Provinces of Nigeria. Bull-. Ent. Res., 15: 1-27. An unidentified reptilian haemogregarine was found in G. tachinoides, The incidence in the fly depended upon the reptilian host: crocodile, frogs, and colubrine snake, 42%; monitors and lizards (Varanus), 457/; and chameleon, 54%. No evidence of pathogenicity to G. tachinoides was found.
Ma, W.-C. & )enlinger, D, L. (1974). Secretory discharge and mieroflora of milk gland in tsetse flies. Nature, 247(5439): 301-303. The results in G. morsitans morsitans which indicated the presence of the posoibly gramnegative bacteria in the lumen of the milk gland were also confirmed by compArative observations of G. austeni Newstead and G. longipalpis pallidiees Atisten. This study indicates some ultrastructural differences between the bacteroids of the mycetome And the bacteria in the milk gland lumen.
274
Notes on some blood parasites collected in Nigeria. Macfie, J. W. S. (1914a). Med. Parasitol., 8: 439-465.
Ann. Trop.
One tsetse fly "out of a number examined" was infected by an unidentified spirochaete. Macfie felt that this spirochaete was distinct from Spirochaeta glossinae described Macfie kept the tsetse in captivity for 22 from G. palpalis by Novy & Knapp (1906). days before it was killed for examination, and the fly did not appear to be affected by the presence of the large number of active spirochaetes.
The occurrence of a spirochaete in the gut of Glossina tachinoides. Macfie, J. W. S. (1914b). The spirochaete was apparently non-pathogenic. Ann. Trop. Med. Parasitol., 8: 464-465.
The results of dissections of tsetse flies at Accra. Macfie, J. W. S. (1915). Lab. Lond., pp. 49-54, 98-99.
Rep. Accra
The Seventy-five flies were dissected and 3 possibly parasitic organisms were found. abdomen of one infected fly was swollen; and it was also hard and pale-coloured ReproDeath was felt to be due, without doubt, to a fungal infection. posteriorly. The hyphae formed duction of the fungus appeared to occur via arthrospore formation. a dense network in which the lower half of the gut and the testicles were firmly embedded. Spirochaetes were found in the gut of another fly and appeared to be morphologically identical to those found by Macfie in the gut of a specimen of G. tachinoides in Nigeria. Also, sporocysts were found in the abdominal cavities of 2 flies and were similar to those found by Chatton and Roubaud (1913) in the abdominal cavity of the same species. Some observations on the seasonal spread of Glossina pallidipes in Moggridge, J. Y. (1936). Italian Somaliland with notes on G. brevipalpis and G. austeni. Bull. Ent. Res., 27: 449466. An unidentified Phycomycete was found in G. brevipalpis. Moggridge noted that the fungus was similar to a pathogenic fungus known in certain areas of Tanzania and that the fungus was present on the abdomen of the flies.
Moloo, S. K. (1972).
Parasitol., 66:
Mermithid parasite of Glossina brevipalpis Newstead.
Ann. Trop. Med.
159.
Two mermithid larvae were found in the coelomic cavity of the one infected adult tsetse. The 5000 flies examined included G. fuscipes, G. brevipalpis and G. pallidipes. Moloo felt that the infection of Glossina was accidental.
The ecology of Glossina morsitans, Westw., and two possible methods Nash, T. A. M. (1933). for its destruction. Part I. Bull. Ent. Res., 24: 107-157. The article includes the statement that a Mr Scott did some work with a fungus and proved Infected larvae had black spots on that it was definitely pathogenic to G. morsitans. their abdomens. Control by parasites and predators of Glossina, pp. 521-532. Nash, T. A. M. (1970). In Mulligan, H. W. (ed.), The African Trypanosomiases, Allen & Unwin Ltd, London. A personal communication from W. H. Potts reports that infection by an unidentified Phycomycete neither prevented completion of development of tsetse larvae nor their deposition.
275 A nematode, tentatively Personal communication to R. F. Myers. Nickle, W. (1973). The identified as Hexamermis sp., was found in the coelomic cavity of G. morsitans. collection was made by S. N. Okiwelu. Three unidentified larval Personal communication to R. F. Myers. Nickle, W. (1974). The collection was made mermithid were found in the coelomic cavities of G. palpalis adults. by K. Riordan.
Investigations on the role of symbionts in tsetse flies (Glossina Nogge, C. (1974). morsitans). Proc. 3rd Int. Cong. Parasitol. (Munich) 2: 947. Attempts were made Symbiont-free flies were obtained by oral treatment with lysozyme. to compensate for the loss of the symbionts by feeding defibrinated blood supplemented by B vitamins. Other techniques such as the raising and lowering of the temperature, the feeding of blood with added sodium oxalate, 7-chlortetracycline, and lysozyme in different concentrations as well as the injection of lysozyme into the coelom resulted in The destruction of the symbionts as well as a decrease in the longevity of the flies. blood meals of these latter flies remained undigested. Novy, F. G. & Knapp, R. E. (1906). J. Infect. Dis., 3: 291-393.
Studies on Spirillum obermeieri and related organisms.
A bacterium, Borrelia glossinae, was described as Spirillum glossinae in 1906 from smears It is listed in Buchanan and Gibbons (Buchanan, R. E. & of flies' stomach contents. 8th Edition. Bergey's Manual of Determinative Bacteriology. 1974. Gibbons, N. E. Baltimore. p. 190) as a Species incertae sedis The Williams and Wilkins Company. because of inadequate description.
The occurrence of intracellular rickettsia-like Pinnock, D. E. & Hess, R. T. (1974). organisms in the tsetse flies, Glossina morsitans, G. fuscipes, G. brevipalpis and Acta Trop., 31: 70-79. G. pallidipes.
Rickettsia-like organisms were observed in electron microscopy preparations the midgut epithelium, including the mycetome, in cells associated with the in developing oocytes. The authors concluded that the presence of a lytic the organisms and that the disruption of cells at the ultrastructural level that the rickettsia-like organisms were parasitic.
of cells of fat body and zone around
suggested
Rapport sur les Travaux de la Mission Scientifique du Katanga Rodhain, J. et al. (1913). (Octobre 1910 a Septembre 1912). Hayez. Bruxelles. 258 p.
Single nematode larvae (possibly Mermis sp.) were found in 4 flies. examined was not given.
The total number
Microorganisms in spermathecae of wild Glossina pallidipes. Rogers, A. (1973). Soc. Trop. Med. Hyg., 67: 299.
Trans. R.
These microorganisms occurred in the lumen of the spermathecae in both inseminated and uninseminated females in wild populations and also occurred at high concentrations in the spermathecae of laboratory-reared females.
Roubaud, E. (1911). 122-132.
Etudes sur les Stomoxydes du Dahomey.
Bull. Soc. Pathol. Exot., 4,
Roubaud attempted to infect by means of direct contact with Entomophthora muscae: 8 G. palpalis, 4 G. tachinoides, and one G. longipalpis and by means of ingestion of the
276
spores: 10 Glossina palpalis. The Glossinas were not infected by either means. The sporangiospores used had been attached to potato leaves and had originated from an infected Stomoxys calcitrans L. However, E. Miller-Kogler (Pilzkrankheiten bei Insekten. Parey. Berlin. 1965, pp. 160-161) has previously pointed out the lack of controls in this study.
Roubaud, E. (1919). Les particularit6s de la nutrition et la vie symbiotique chez les mouches ts6tses. Ann. Inst. Pasteur, 33: 489-536.
The mycetome is located Cicadomyces sp. is a symbiote of G. palpalis and G. tachinoides. in the middle of the anterior segment of the mid-gut, where absorption but no digestion occurs (Wigglesworth, V. B. (1972). The Principles of Insect Physiology. Chapman & Hall. London, p. 523).
Roubaud, E. & Treillard, M. (1935). Un coccobacille pathogene pour les mouches Tsetses. C. R. Hebd. Seances Acad. Sci. (Paris) 201: 304-306. This microorganism, Bacterium mathisi, which initially showed promise as a biocontrol agent has neither been maintained in culture nor is it a recognized valid taxon. Roubaud & Treillard (1935) found that the bacterium was taken in via the proboscis during feeding on guinea pigs whose hair and skin had been coated with the bacterium. The flies are apparently killed by a septicemic infection within the coelom within 1-8 days. Several insects were shown to be susceptible when inoculated with bacterial suspensions; however, Culex pipiens, Stegomyia sp. and Aedes aegypti survived ingestion of the microbe.
Roubaud, E. & Treillard, M. (1936). Infection experimentale de Glossina palpalis par un coccobacille pathogane pour les muscides. Bull. Soc. Pathol. Exot., 29: 145-147. No naturally occurring infection by Bacterium mathisi was found in the several hundred Glossina palpalis shipped from Uganda in the pupal stage. The low number of flies used in the experiments makes interpretation difficult; however, it should be noted that a septicemia was not indicated in all cases of adult mortality. When G. morsitans from Tanzania were used in experiments, they died of a septicemia in 1 to 3 days after feeding on contaminated guinea pigs.
Swynnerton, C. F. M. (1936). The tsetse flies of East Africa. A first study of their ecology, with a view to their control. Trans. R. Entomol. Soc. Lond., 84: 1-580. An unidentified Phycomycete was found infecting G. palpalis. Material for the cultivation of the fungus was sent to Johannesburg; however, the study does not appear to have been continued. Swynnerton felt that the hope of success using this particular fungus was small because it seemed to be fatal only during periods of wet weather.
Thomson, W. E. F. (1947).
Nematodes in tsetse.
Ann. Trop. Med. Parasitol., 41:
164.
The nematodes were located in the coelomic cavity and had an average length of 79 mm. InterVago, C. & Meynadier, S. pp. 24, 27, In Strains of Entomophagous Microorganisms. WPRS Bulletin national Organization for Biological Control of Noxious Animals and Plants.
1973/3. Aspergillus flavipes and Penicillium janthinellum were isolated from G. fusca congolensis in 1965. Both fungi have been previously isolated from soil in Africa.
277 Vanderyst, P. H. (1923). 32-42.
La prophylaxie contre la trypanose humaine.
Rev. Med. Angola, 4:
A local fly "Nkulu nyansi" was very rapidly infected with Entomophthora muscae when placed in a tube with one of the infected M. domestica L. received from Europe; however, The experiment was later G. palpalis was not infected under the same conditions. supplemented by adding locally obtained M. domestica, and these were also not infected. One of the tsetses used did die during the course of the experiment but did not show any signs of a disease. The infection of the fly "Nkulu nyansill was felt to be enhanced by Other experiments with this local fly gave negative injury during insect procurement. results. This is the first attempt at assessing the virulence of a microbial pathogen derived from outside Africa. La transmission cyclique de races r6sistantes de van Hoof, L. & Henrard, C. (1934). Ann. Soc. Belge Med. Trop., 14: 109-144. Trypanosoma gambiense par Glossina palpalis.
Hyphae of an unidentified fungus occurred in the coelomic cavity, in the gut, in the musculature of the thorax and in the proventriculus of G. palpalis. These authors cite a reference (Lloyd & Johnson, 1922), in the text to a similar infection but do not include it in the bibliography. Etudes sur Recherches sur les champignons pathogenes pour les glossines. Vey, A. (1971). Rev. Elev. Med. Glossina fusca congolensis Newst. et Evans. en Republique Centrafricaine. Vet. Pays Trop. (N.S.) 24: 577-579. Two fungal species were isolated from tsetse pupae, Absidia repens and Penicillium Both are common soil inhabitants. Experimental infections of tsetse larvae lilacinum. are reported. Larvae were wounded prior to incubation with the spore suspensions.
An unidentified coccoid bacterium was Personal communication to R. Nolan. Vey, A. (1974). isolated from Glossina pupae at the Laboratory I.E.M.U.T. de Farcha (Chad). Aspergillus ochraceus was isolated from Glossina pupae into pure culture in the Laboratory I.E.M.U.T. de Farcha (Chad). A. ochraceus has also been found in material of Theobroma cacao from the Gold Coast (Dade, H. A. 1940. A revised list of Gold Coast Fungi and Plant Diseases. Bulletin of Miscellaneous Information No. 6, pp. 205Kew Bulletin. Royal Botanic Gardens.
247). Wallace, J. M. (1931). Parasitol., 25: 1-19.
Micro-organisms in the gut of Glossina palpalis.
Ann. Trop. Med.
A fairly thorough study which would indicate that G. palpalis does not readily pick up bacteria from contaminated surroundings, such as fly boxes; however, during feeding Wallace felt that some earlier workers experiments organisms can be introduced into the gut. had introduced bacteria which were surface contaminants into gut preparations during He also observed that in every instance where Trypanosoma grayi was present the dissection. This latter finding is contrary to those of Carpenter (1912). bacteroids were also present. Wallace's comments with regard to the general inability of G. palpalis to acquire bacteria within the gut by feeding through contaminated skin or hair because of a possible downward flow of saliva or because saliva is bactericidal is contrary to the findings of Roubaud & Treillard (1935) using G. morsitans.
