Progress 07/01/00 to 06/30/05
Outputs
Forty-five new species of black flies were described from North America and one new species was described from Australia. Identification keys were provided for the larvae, pupae, males, and females of all 255 known species of black flies in North America, and all bionomic and economic information for each species was compiled. Chromosomal analyses were presented for many species in North America, South America, and Europe. The Holarctic black fly Simulium noelleri, for example, was shown cytogenetically to consist of three distinct cytotypes, possibly representing different species. Sibling species in the Simulium vittatum complex were differentiated molecularly based on random amplified polymorphic DNA. An analysis of the cytogenetics of a laboratory colony of black flies indicated that all sex chromosomes and autosomal polymorphisms in the source population were still present in the colony after nearly two decades. The study also demonstrated a probable example of
pseudo-partial sex linkage involving X-linked sequences. Faunal surveys provided insight into the species composition of black flies in selected areas of the world such as the Great Smoky Mountains National Park (Tennessee), Guadalupe Mountains National Park (Texas), Kansas, Central Amazonia (Brazil), Morocco, Prince Edward Island (Canada), and Sachsen-Anhalt State (Germany). Ecological investigations demonstrated that black flies play an important role in ecological processes in the boreal region; for example, they are a major component of the diet of some carnivorous plants. Investigations of symbiotes of black flies resulted in a morphological and molecular description of one new species of microsporidium and a phylogeny of the microsporidian genus Caudospora. Studies of trichomycete fungi associated with the digestive tracts of larval black flies showed that these fungi have seasonal trends in prevalence and that ecological conditions can serve as predictors of their prevalence in
streams.
Impacts
Combined chromosomal and morphological investigations provide powerful insights into the black flies of the world, demonstrating similarities between the faunas of the Old and New Worlds and allowing previously independent or seemingly unrelated information to be used to solve pest problems. This approach has made the black fly fauna of North America one of the taxonomically best known groups of insects in all life stages. Because sound taxonomy is the spine of pest management and basic biological research, improvements can continue to be made in the control of black flies while ecological investigations can use black flies as model organisms. The concomitant study of black fly symbiotes, such as trichomycete fungi, promises to open new possibilities for using natural enemies in the biological control of pest species.
Publications
- Adler, P. H. 2004. Polytene chromosomes. In J. L. Capinera (ed.). Encyclopedia of Entomology. Kluwer Academic Publishers.
- Adler, P. H. 2004. Sibling species. In J. L. Capinera (ed.). Encyclopedia of Entomology. Kluwer Academic Publishers.
- Adler, P. H. 2004. Black flies. In J. L. Capinera (ed.). Encyclopedia of Entomology. Kluwer Academic Publishers.
- Adler, P. H. and Wills, W. 2003. Legacy of death: the history of arthropod-borne human diseases in South Carolina. American Entomologist 49: 216-228.
- Alvan-Aguilar, M. A., N. Hamada, Adler, P. H and S. L. Bessa Luz. 2005. Cytotaxonomy of Simulium cauchense Floch and Abonnenc and Simulium quadrifidum Lutz (Diptera: Simuliidae) in Brazilian Amazonia. Memorias do Instituto Oswaldo Cruz 100: 249-257.
- Malmqvist, B., Adler, P. H and Strasevicius, D. 2004. Testing hypotheses on egg number and size in blackflies (Diptera: Simuliidae). Journal of Vector Ecology 29: 248-256.
- McCreadie, J. W., Adler, P. H and N. Hamada. 2005. Patterns of species richness for blackflies (Diptera: Simuliidae) in the Nearctic and Neotropical regions. Ecological Entomology 30: 201-209.
- Minhas, M. S., C. L. Brockhouse and Adler, P. H. 2005. The black fly (Diptera: Simuliidae) fauna of Prince Edward Island, Canada. Northeastern Naturalist 12: 67-76.
- Reeves, W. K. and Adler, P. H. 2004. Black flies (Diptera: Simuliidae) and their symbiotes from Guadalupe Mountains National Park, Texas. Studia Dipterologica 11: 3-7.
- Werner, D. and Adler, P. H. 2005. A faunistic review of the black flies (Simuliidae, Diptera) of the federal state of Sachsen-Anhalt, Germany. Abhandlungen und Berichte fur Naturkunde 27: 205-245.
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Progress 01/01/04 to 12/31/04
Outputs
Forty-three new species of North American black flies and one new species of Australian black fly were described and illustrated. All knowledge of black flies in North America was summarized in the publication of an illustrated book entitled the "Black Flies (Simuliidae) of North America," which includes identification keys, bionomic entries and distribution maps for each of the 255 known species, phylogenetic reconstructions, cytological and morphological summaries, economic impact assessments, control strategies, and basic behavioral and ecological information. Users will now be able to identify black flies throughout the continent and to access all information known about each species. Collaborative molecular work provided supporting evidence that the two morphologically indistinguishable members of the Simulium vittatum complex are valid species. Additional collaborative molecular efforts demonstrated that vertebrate blood meals from the guts of wild-caught black
flies could be analyzed, based on the cytochrome b gene, to determine the precise hosts of different species of black flies. Black flies in areas such as the boreal biome are so abundant that they drive ecosystem processes, both as larvae and as adults. In fact, an unusual new form of predation on black flies was documented, with the discovery that black flies are so abundant that they constitute the dominant prey of some boreal populations of streamside insectivorous plants of the genus Pinguicula. A large dataset of cytologically analyzed black flies from the Nearctic and Neotropical Regions showed that as regional richness increased, local diversity reached an asymptote in which further increases in regional richness were not matched by increases in local richness, suggesting that simuliid communities are saturated, which is consistent with the current view of lotic communities as nonequilibrium systems. Species richness of black flies was found to be no greater in tropical than in
temperate streams.
Impacts
Combined chromosomal and morphological investigations continue to provide new insights into the black flies of the world, demonstrating similarities between the faunas of the Old and New Worlds and allowing previously independent or seemingly unrelated information to be used in solving problems. As a result of our ongoing survey, the black fly fauna of North America is now one of the taxonomically best known groups of insects in all life stages. Because sound taxonomy is the spine of pest management and solid biological research, improvements can continue to be made in the control of black flies while ecological investigations can use black flies as model organisms. The concomitant study of black fly symbiotes, such as trichomycete fungi, promises to open new possibilities for using natural enemies in the biological control of pest species.
Publications
- Adler, P. H. 2004. Black flies, the Simuliidae. Pp. 127-140. In W. C. Marquardt (ed.). Biology of Disease Vectors, 2nd edition. Elsevier Academic Press, San Diego, CA. 785 pp. [issued December 2004].
- Adler, P. H. & B. Malmqvist. 2004. Predation on black flies (Diptera: Simuliidae) by the carnivorous plant Pinguicula vulgaris (Lentibulariaceae) in northern Sweden. Entomologica Fennica 15: 124-128.
- Adler, P. H., D. C. Currie & D. M. Wood. 2004. The Black Flies (Simuliidae) of North America. Cornell University Press, Ithaca, NY. xv + 941 pp. + 24 color plates.
- Duncan, G., P. H. Adler, K. P. Pruess & T. O. Powers. 2004. Molecular differentiation of two sibling species of the black fly Simulium vittatum (Diptera: Simuliidae) based on random amplified polymorphic DNA. Genome 47: 373-379.
- Malmqvist, B., P. H. Adler, K. Kuusela, R. W. Merritt & R. S. Wotton. 2004. Black flies in the boreal biome, key organisms in both terrestrial and aquatic environments: a review. Ecoscience 11: 187-200.
- Malmqvist, B., D. Strasevicius, O. Hellberg, P. H. Adler & S. Bensch. 2004. Vertebrate host specificity of wild-caught blackflies revealed by mitochondrial DNA in blood. Proceedings of the Royal Society of London B (Supplement), Biology Letters 271: S152-S155.
- Moulton, J. K., P. H. Adler & J. Prince. 2004. An unusual new species of Paracnephia Rubtsov (Diptera: Simuliidae) from Western Australia. Zootaxa 409: 1-12.
- Reeves, W. K., P. H. Adler, W. L. Grogan, Jr. & P. E. Super. 2004. Hematophagous and parasitic Diptera (Insecta) in the Great Smoky Mountains National Park, USA. Zootaxa 483: 1-44.
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Progress 01/01/03 to 12/31/03
Outputs
Investigations of the systematics and natural enemies of the black flies of North America, in the context of the world fauna, have continued. Surveys of the southeastern and far northern portions of the continent revealed one species previously unknown beyond Scandinavia. Simulium annulitarse was discovered breeding in small streams on the western shore of Hudson Bay, bringing the total number of known North American species to 255 and further illustrating the intimate connection between the Nearctic and Palearctic faunas. Populations of this species in the two zoogeographic regions are virtually identical chromosomally. Thirty-three (13%) of the species in North America are now known to be shared with the Palearctic Region. The larva and pupa of a new species of black fly from Nova Scotia, Canada, were described and illustrated. This new species, Simulium rothfelsi, is most closely related to S. johannseni of midwestern North America and to a new species from the
southeastern piedmont ecoregion. It can be distinguished from all species in North America by the unique pupal gill, which bears two swollen dorsal filaments and two slender ventral filaments. The new species passes the winter as eggs and completes its larval and pupal development in a few weeks during May. To gain insight into relationships of black flies and the regulation of their populations, investigations of their symbiotes, particularly trichomycete fungi, were undertaken. Larval black flies from more than 60 streams in South Carolina were surveyed for the midgut-inhabiting trichomycete fungus Harpella melusinae. Our results indicate that nearly all species of black flies are colonized by this fungus, and that certain ecological conditions, such as stream pH, can serve as predictors of the prevalence of the fungus. Because much of the taxonomy of trichomycetes is based on zygospores, methods were developed to induce their production, using an application of pH 10 potassium
hydroxide in culture media. This procedure allowed the first description of the zygospores of additional trichomycete species.
Impacts
Combined chromosomal and morphological investigations continue to provide new insights into the black flies of the world, demonstrating similarities between the faunas of the Old and New Worlds and allowing previously independent or seemingly unrelated information to be used in solving problems. As a result of our ongoing survey, the black fly fauna of North America is now one of the taxonomically best known groups of insects in all life stages. Because sound taxonomy is the spine of pest management and solid biological research, improvements can continue to be made in the control of black flies while ecological investigations can use black flies as model organisms. The concomitant study of black fly symbiotes, such as trichomycete fungi, promises to open new possibilities for using natural enemies in the biological control of pest species.
Publications
- Adler, P. H., C. L. Brockhouse and D. C. Currie. 2003. A new species of black fly (Diptera: Simuliidae) from Nova Scotia. Proceedings of the Entomological Society of Washington 105: 9- 12.
- Beard, C. E. and P. H. Adler. 2003. Zygospores of selected Trichomycetes in larval Diptera of the families Chironomidae and Simuliidae. Mycologia 95: 317-320.
- Beard, C. E., J. W. McCreadie and P. H. Adler. 2003. Prevalence of the trichomycete fungus Harpella melusinae (Harpellales: Harpellaceae) in larval black flies (Diptera: Simuliidae) across a heterogeneous environment. Mycologia 95: 577-583.
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Progress 01/01/02 to 12/31/02
Outputs
A combined cytogenetic-morphological-ecological approach was used to investigate selected black flies of the world, with an emphasis on revealing sibling species and understanding ecological relationships. One new species, in the subgenus Psilopelmia, was named and described from New Mexico. The black flies of Kansas were surveyed and their pest status over the past three decades was documented. Fourteen species were recorded from Kansas, including four new state records and two major pests, namely Simulium meridionale and Simulium vittatum. The death in Kansas of a double yellowhead Amazon parrot from black fly attacks represents the first documented record of North American black flies causing economic loss to owners of exotic birds other than ratites. Similarly, the blood-feeding habits of five species of black flies on black grouse in Finland were documented, and included the first host records for several species. Efforts were made to predict species
distributions based on habitat characteristics of larvae in the Central Amazon River Basin. Species occurrence was highly predictable, ranging from about 80 to 90% for the six most common species. Stream size and the presence of impoundments were the most important predictors of species distributions, reinforcing similar discoveries that we have made in other regions of the world. Additional ecological work was aimed at the symbiotic trichomycete fungi that inhabit larval black fly guts. A study of the seasonality of eight species of trichomycetes that colonize South Carolina black flies revealed that more than 93% of larvae harbored at least one species of trichomycete and that fungal prevalence differed significantly with season. The chromosomes of four species of black flies, two from the United States and two from Brazil, were described in detail. Of particular interest was a cytogenetic study of changes in laboratory colonies of Simulium vittatum cytospecies IS-7 over an 18-year
period. All sex chromosomes and major autosomal polymorphisms that were present in the source population were still represented in the laboratory colonies. However, the extent of sex linkage and the frequencies of four of the five major autosomal inversions had changed significantly in at least one colony, possibly because of bottlenecks experienced by the colony. The absence of males homozygous for the IS-7 inversion in both field and colony material was explained by postulating that the Y2 chromosome is entirely absent or acts as a rare reproductive lethal. This example possibly represents pseudo-partial sex linkage involving the X-linked sequences.
Impacts
The study of black flies, the second most important group of medically important insects, from a world perspective, using multiple investigative approaches, provides the greatest insight into problems of species identification and relationships with the environment. The discovery of new species and universal biological characteristics of black flies, including predictors of species distributions, will greatly enhance efforts to control the pest species.
Publications
- Brockhouse, C. L. & P. H. Adler. 2002. Cytogenetics of laboratory colonies of Simulium vittatumcytospecies IS-7 (Diptera: Simuliidae). Journal of Medical Entomology 39: 293-297.
- Hamada, N., J. W. McCreadie & P. H. Adler. 2002. Species richness and spatial distributions of blackflies (Diptera: Simuliidae) in streams of Central Amazonia, Brazil. Freshwater Biology 47: 31-40.
- Mock, D. E. & P. H. Adler. 2002. Black flies (Diptera: Simuliidae) of Kansas: review, new records, and pest status. Journal of the Kansas Entomological Society 75: 203-213.
- Moulton, J. K. & P. H. Adler. 2002. Taxonomy and biology of Simulium clarkei Stone & Snoddy (Diptera: Simuliiidae), a poorly known black fly of the southeastern United States. Zootaxa 31: 1-7.
- Moulton, J. K. & P. H. Adler. 2002. A new species of Simulium (Psilopelmia) Enderlein (Diptera: Simuliidae) from southern New Mexico, USA. Studia Dipterologica 9:213-218.
- Ojanen, U., O. Ratti. P. H. Adler, K. Kuusela, B. Malmqvist & P. Helle. 2002. Blood feeding by black flies (Diptera: Simuliidae) on the black grouse (Tetrao tetrix) in Finland. Entomologica Fennica 13: 153-158.
- Rios-Velasquez, C., N. Hamada, & P. H. Adler. 2002. Cytotaxonomy of Simulium goeldii Cerqueira & Nunes De Mello and Simulium ulyssesi (Py-Daniel & Coscaron) (Diptera: Simuliidae) in Central Amazonia, Brazil. Journal of Insect Systematics and Evolution 33: 113- 120.
- Adler, P. H. & J. W. McCreadie. 2002. Black flies (Simuliidae). Pp. 185-202. In G. R. Mullen and L. A. Durden (eds.). Textbook of Medical and Veterinary Entomology. Academic Press, New York.
- Beard, C. E. & P. H. Adler. 2002. Seasonality of trichomycetes in larval black flies from South Carolina, USA. Mycologia 94: 200-209.
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Progress 01/01/01 to 12/31/01
Outputs
Progress toward a global understanding of black flies was made by cytogenetically and morphologically investigating the relationships of North American and European black flies. One of the world's most widespread black flies, Simulium noelleri, was analyzed chromosomally and morphologically. Reproductive isolation of S. noelleri from the morphologically identical, Nearctic species, S. decorum, was demonstrated. Simulium noelleri was determined to be a single species composed of at least three distinct cytotypes differing mainly in their sex chromosomes. Cytotype A, representing the type species of S. noelleri, is found in Germany, England, and western Russia. Cytotype B occurs in Sweden and western Canada, indicating the close association between the northern Nearctic and Palearctic simuliid faunas. Cytotype C is known only from Armenia. The Nearctic name corbis and Palearctic name relictum were synonymized with murmanum. Thus, S. murmanum has a Holarctic
distribution. Three species in the Prosimulium hirtipes group in Morocco were investigated chromosomally. Each species (P. latimucro, P. rufipes, and P. tomosvaryi) is chromosomally unique; one or more of these Moroccan species is probably distinct from the European species of the same names. An annotated summary of all species known from Morocco was prepared as an aid to understanding the distribution of black flies at the southern extreme of the Palearctic Region.
Impacts
Successful management of black flies depends on a sound taxonomic foundation that permits precise identification and targeting of the actual pest species. Only through increased taxonomic rigor can more effective and economical management efforts be achieved. The demonstration that numerous species are common to both the Nearctic and Palearctic Regions will aid the global synthesis of biological information on these species, and will allow the efforts aimed independently at pest species in each zoogeographic area to be directed against the total pest problem.
Publications
- Evans, C. L. and P. H. Adler. 2000. Microsculpture and phylogenetic significance of the spermatheca of black flies (Diptera: Simuliidae). Canadian Journal of Zoology 78: 1468-1482.
- Raastad, J. E. and P. H. Adler. 2001. The identity of Simulium murmanum Enderlein, 1935 (Diptera: Simuliidae). Aquatic Insects 23: 183-186.
- Adler, P. H. and B. Belqat. 2001. Cytotaxonomy of the Prosimulium hirtipes species group (Diptera: Simuliidae) in Morocco. Journal of Insect Systematics and Evolution 32: 411-418.
- Adler, P. H. and E. A. Kachvoryan. 2001. Cytogenetics of the Holarctic Black Fly Simulium noelleri (Diptera: Simuliidae). Canadian Journal of Zoology 79: 1972-1979.
- Hamada, N. and P. H. Adler. 2001. Bionomia e chave para imaturos e adultos de Simulium(Diptera: Simuliidae) na Amazonia Central, Brasil. Acta Amazonica 31: 109-132.
- Belqat, B., P. Adler and M. Dakki. 2001. Distribution summary of the Simuliidae of Morocco with new data for the Rif Mountains. British Simuliid Group Bulletin 17: 10-16.
- Adler, P. H., R. W. Merritt, J. F. Burger and D. P. Molloy. 2001. A brief history of Northeast Regional Project NE-118 in the USA. British Simuliid Group Bulletin 17: 16-20.
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Progress 01/01/00 to 12/31/00
Outputs
Simuliid pathogens, especially microsporidia and trichomycete fungi, were investigated as possible indicators of species limits and phylogenetic relationships, as well as for their potential in the biological control of black flies. A new species of caudosporid microsporidium was described ultrastructurally and molecularly from the eastern United States, where it was especially conspicuous in the coastal plain. The highest densities of infected simuliid larvae ever recorded were found in the type host (Cnephia ornithophilia) in South Carolina. This study of microsporidia represents the first molecular characterization of a simuliid microsporidium and the first hypothesis of caudosporid relationships based on the phylogenetic relationships of the hosts. The caudosporid microsporidia are more host specific than most other species of microsporidia currently known in black flies. New records of trichomycete fungi were found in black flies in South Carolina and
similarities in the trichomycete fungi of black flies throughout the Holarctic Region was demonstrated. Certain trichomycete characters (e.g., spore size) currently used in species diagnoses were found to vary significantly depending on the host and medium in which they grew, casting doubt on some of the current species concepts of trichomycetes. The investigated trichomycetes were common in numerous species of black flies and did not appear to be host specific.
Impacts
Biological control of black flies depends on a sound taxonomy of both the pest species and their pathogens. The discovery of new species of pathogens in black flies, and their molecular, structural, and host charcterization, will provide the basis for using new biological control agents in the management of black flies.
Publications
- Adler, P. H., J. J. Becnel & B. Moser. 2000. Molecular characterization and taxonomy of a new species of Caudosporidae (Microsporidia) from black flies (Diptera: Simuliidae), with host- derived relationships of the North American caudosporids. Journal of Invertebrate Pathology 75: 133-143.
- Beard, C. E. & P. H. Adler. 2000. Bionomics, axenic culture, and substrate-related variation in trichospores of Smittium megazygosporum. Mycologia 92: 296-300.
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