Some boundary layer characteristics of microhabitats occupied by larval black flies (Diptera: Simuliidae)

Microhabitats occupied by larvae of 12 species or complexes of black flies at 30 sites in western Canada were investigated for fluid flow parameters. An attached boundary layer was a feature of almost all microhabitats occupied by larvae of black flies. Although not measured directly, acceleration of water resulting from flow around an object appeared to alter flow characteristics and make microhabitats more favourable. This predictability in the flow characteristics of microhabitats occupied by larvae of black flies implies that the contagious dispersion of these larvae is a response to the heterogeneity of flow over a stream bed. It is suggested that microhabitats characterized by suitable flow characteristics may constitute a resource that limits population size. Larvae occupied microhabitats with both laminar and turbulent boundary layers, and were found in microhabitats with a wide range of water velocities. Mainstream flow parameters investigated were; water velocity, water depth, and whether flow was subcritical, critical or supercritical. Boundary layer parameters investigated were; whether flow was laminar or turbulent, and the point at which separation from the substratum occurred.     

Context-dependent symbiosis between black flies (Diptera: Simuliidae) and trichomycete fungi (Harpellales: Legeriomycetaceae)

The context-dependent nature of a symbiotic relationship between a trichomycete fungus ( Smittium culisetae ) and a larval black fly ( Simulium vittatum ) is demonstrated in the present study. No significant difference was found between the size of larvae colonized by trichomycetes and those free of trichomycetes, regardless of the trichospore dosage or initial age of the larvae. This trend suggests that the trichomycete has no detectable effect on host fitness, indicating a commensalistic relationship. However, in half of the experiments, stressed (i.e. starved) larvae exposed to trichospores at a dosage of 20 000 spores ml⁻¹ had significantly higher survival than did trichomycete-free larvae, indicating a mutualistic relationship. Trichomycetes in adult female black flies can replace the ovaries. The symbiotic association between trichomycetes and simuliids, therefore, is dynamic: commensalistic when larvae are well fed, mutualistic when larvae are starved, and parasitic in adults. The trichomycete-black fly relationship represents a rare case of symbiosis shifting among three states.     

Seasonal occurrence of black flies (Diptera: Simuliidae) in a desert stream receiving trout farm effluent

The distribution and abundance of black flies (Diptera: Simuliidae) in a small desert stream were influenced by environmental changes caused by recharge of water supply storage basins and an aquaculture operation. Simulium virgatum was the most abundant benthic insect collected in Whitewater Canyon (Riverside County, CA) after April; however, it was never found in trout farm effluent where the ammonium‐nitrogen concentration was > 0.25 mg/liter. S. virgatum densities downstream of the input of water from the Colorado River aqueduct were lower than at other sampling sites in the Whitewater River. Simulium tescorum, an especially anthropophilic black fly, was most abundant during February and March, was not collected from late spring through early autumn, and was found only in the highly enriched, less variable flow of trout farm effluent. The mean concentrations of ammonium nitrogen and nitrate nitrogen in the trout farm effluent nearly 1 km from the fish holding ponds were ten and two times, respectively, the ambient levels in the Whitewater River upstream of the effluent discharge point. A combination of factors probably contributed to the presence of S. tescorum in the trout farm effluent including homogenization of the flow regime, enrichment of larval resources, and the development of riparian vegetation that provided oviposition and attachment sites.     

Potentialities of mermithid nematodes for the biocontrol of blackflies (Diptera: Simuliidae)--a review

Mermithids comprise a family of nematodes which invariably kill/sterilize their insect host(s). These nematodes have considerable potential as biocontrol agents of agricultural insect pests and medically important insect vectors. More specifically, mermithid nematodes appear to regulate natural population of blackflies. The taxonomy of the Mermithidae has been only partially evaluated and the taxonomic status of many representatives is uncertain. At least three mermithid genera and species parasitize North American blackflies, although a more varied mermithid fauna probably exists. The host specificity of mermithid parasites of simuliids is variable, but these nematodes do not appear to infect other stream fauna. The sporadic distribution of mermithid parasites of simuliids among potential biotopes may be associated with a relatively inefficient mode of dispersal for such nematodes. Detailed information is lacking concerning stages in the life cycles of these pathogens and their synchronization with the simuliid host. Mermithids cause pathogenic effects upon several blackfly tissues, although no information is available concerning physiological manifestations of mermithid parasitism in blackflies. A brief review of the present state of knowledge of simuliid taxonomy and bionomics is presented. The physiology of blackflies and their mermithid parasites has been largely ignored. The potentialities of mermithid nematodes for the biocontrol of blackflies are assessed and a feasible research programme presented, in relation to the present state of knowledge of mermithid-simuliid interrelationships and related areas of insect nematology.     

Global diversity of black flies (Diptera: Simuliidae) in freshwater

Black flies are a worldwide family of nematocerous Diptera whose immature stages are confined to running waters. They are key organisms in both aquatic and terrestrial ecosystems, but are perhaps best known for the bloodsucking habits of adult females. Attacks by black flies are responsible for reduced tourism, deaths in wild and domestic birds and mammals, and transmission of parasitic diseases to hosts, including humans. About 2,000 nominal species are currently recognized; however, certain geographical regions remain inadequately surveyed. Furthermore, studies of the giant polytene chromosomes of larvae reveal that many morphologically recognized species actually consist of two or more structurally indistinguishable (yet reproductively isolated) sibling species. Calculations derived from the best-known regional fauna—the Nearctic Region—reveal that the actual number of World black fly species exceeds 3,000. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: E.V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Digestive system of larval black flies (Diptera: Simuliidae): structure and phylogeny

Abstract. Larval gut characters of 45 species of black flies, representing 17 subgenera and 15 genera, were studied for their utility in taxon diagnosis and phylogenetic inference. Phase-contrast and scanning electron microscopy revealed a wealth of new characters in the esophageal armature of the foregut. Larval gut morphology, particularly characters of the esophageal armature, varied significantly among taxa, permitting generic diagnoses. A cladogram inferred from 13 gut characters supported a sister-group relationship between Parasimulium and the remaining simuliids, but did not show a strong phylogenetic signal for the current, well-supported concept of tribal level monophyly.     

Scale, time, space, and predictability: species distributions of preimaginal black flies (Diptera: Simuliidae)

This study examines species-specific distributions of preimaginal black flies (Diptera: Simuliidae) over two large spatial scales (within and across ecoregions) and two seasons (spring and summer). The study area in the western two-thirds of South Carolina, United States, was divided into three ecoregions (Mountains, Piedmont, Sandhills). The mean correct classification of species distributions among stream sites was 81.3%. Predictability of species distributions varied with spatial scale, location, and time. Species occurrence was not independent of ecoregion; distributions of 12 of the 13 species examined at this scale were predictable on the basis of changes in water column (temperature, percentage dissolved oxygen) and channel characters (velocity, streambed-particle size). However, with the exception of the Mountains during the summer, predicting species distributions within ecoregions was far less successful than predicting distributions across these regions; predictability was particularly poor in the Sandhills. Seasonal effects on predictability were most pronounced in the Mountains. As stream sites became more homogeneous, predictive capability declined. Species-level identifications, aided by cytogenetic techniques, were of paramount importance in recognizing the association between species distributions and stream conditions, emphasizing that considerable ecological information is lost when species are not considered. This study emphasizes the critical role of taxonomic resolution, linked with spatial and temporal scale of observation, in elucidating patterns of species assemblages. It also demonstrates that predictability of species distributions at a fixed scale can vary with geographic location and time. Received: 2 May 1997 / Accepted: 22 September 1997     

Insular black files (Diptera: Simuliidae) of North America: tests of colonization hypotheses

Aim To understand factors that facilitate insular colonization by black flies, we tested six hypotheses related to life‐history traits, phylogeny, symbiotes, island area, and distance from source areas. Location Four northern islands, all within 150 km of the North American mainland, were included in the study: Isle Royale, Magdalen Islands, Prince Edward Island, and Queen Charlotte Islands. Methods Immature black flies and their symbiotes were surveyed in streams on the Magdalen Islands, and the results combined with data from similar surveys on Isle Royale, Prince Edward Island, and the Queen Charlotte Islands. Black flies were analysed chromosomally to ensure that all sibling species were revealed. Tests of independence were used to examine the frequency of life‐history traits and generic representation of black flies on islands vs. source areas. Results A total of 13–20 species was found on each of the islands, but no species was unique to any of the islands. The simuliid faunas of the islands reflected the composition of their source areas in aspects of voltinism (univoltine vs. multivoltine), blood feeding (ornithophily vs. mammalophily), and phylogeny (genus Simulium vs. other genera). Five symbiotic species were found on the most distant island group, the Magdalen Islands, supporting the hypothesis that obligate symbiotes are effectively transported to near‐mainland islands. An inverse relationship existed between the number of species per island and distance from the source. The Queen Charlotte Islands did not conform to the species–area relationship. Main conclusions The lack of precinctive insular species and an absence of life‐history and phylogenetic characteristics related to the presence of black flies on these islands argue for gene flow and dispersal capabilities of black flies over open waters, possibly aided by winds. However, the high frequency of precinctive species on islands 500 km or more from the nearest mainland indicates that at some distance beyond 100 km, open water provides a significant barrier to colonization and gene exchange. An inverse relationship between number of species and distance from the source suggests that as long as suitable habitat is present, distance plays an important role in colonization. Failure of the Queen Charlotte Islands to conform to an area–richness trend suggests that not all resident species have been found.     

Prevalence of the trichomycete fungus Harpella melusinae (Harpellales: Harpellaceae) in larval black flies (Diptera: Simuliidae) across a heterogeneous environment

A total of 2063 mid- to late-instar larval black flies were collected from 64 stream sites in South Carolina and screened for the presence of the trichomycete fungus Harpella melusinae. Sixteen of 18 host species were colonized by H. melusinae on at least one occasion. Prevalence of H. melusinae in larvae of Simulium tuberosum cytospecies "A" was highest in acidic streams with low conductivity, whereas H. melusinae colonized larvae of Simulium verecundum most frequently in slower-moving streams. Ecological conditions, therefore, can serve as predictors of the prevalence of H. melusinae. Prevalence in host larvae was significantly lower in the Piedmont ecoregion than in the Mountain ecoregion. Prevalence did not differ in the host species S. verecundum across ecoregions, suggesting that different prevalences among host species might indicate some host preference. The prevalence of H. melusinae differed significantly between two univoltine host species (Simulium venustum and Prosimulium magnum) at the same site but not between two multivoltine host species (S. tuberosum cytospecies "FG" and S. tuberosum cytospecies "CDE"), suggesting that host life history could be important in determining fungal prevalence.     

Long-term emergence patterns of black flies (Diptera: Simuliidae) in northwestern Pennsylvania

Spatial and temporal patterns of black fly emergence at three sites over a 7-year to 11-year period are presented. This study is the longest, continuous collection of emerging black flies ever undertaken. Because ovipositing females were able to enter emergence traps, analysis of emergence patterns for the Simulium vittatum Zetterstedt and S. tuberosum Lundström complexes, was restricted to males. Year, season and stream temperature had significant effects on mean emergence for these complexes. Sampling stations on the same system and within close proximity exhibited synchronous emergence patterns, whereas streams in close proximity but in separate basins showed asynchronous patterns. Dissimilarities might reflect divergent temperature regimes. We suggest that emergence-trap data might provide a less variable picture of community structure than larval data.     

A molecular revision of the taxonomic status of mermithid parasites of black flies from Quebec (Canada)

The four currently recognized mermithid (Nematoda) species parasitizing black flies (Diptera: Simuliidae) from Northeast America were distinguished using discriminatory PCR primers aimed at COI and 18S rDNA. Isomermis wisconsinensis, Gastromermis viridis and Mesomermis camdenensis were easily differentiated using either genomic target, even for juvenile mermithids damaged beyond morphological recognition. However, specimens from Mesomermis flumenalis being identical in external morphology and producing a unique-sized PCR product were classified by sequence data into four clearly distinguished molecular variants. This quartet was made of two winter and two summer ‘physiological variants’, including one which also belonged to, but diverged early from the rest of the Mesomermis genus. Combining the multiplex PCR and sequencing approaches allowed for the characterization of a multiple parasitism which simultaneously implicated I. wisconsinensis and two M. flumenalis variants. With another instance where parasites were identified by morphology only, this is the first report of black fly parasitism by multiple mermithid species. A phylogenetic tree built by combining our sequences to previous GenBank entries likely indicates a monophyletic origin for the mermithid family, but also suggests that differentiation between parasite genera sometimes occurred before the evolutionary emergence of the actual host group.     

Molecular phylogeny of black flies (Diptera: Simuliidae) from Thailand, using ITS2 rDNA

The sequences of the second internal transcribed spacer (ITS2) of ribosomal DNA (rDNA) were determined for 40 black fly species from Thailand, belonging to 4 subgenera of the genus Simulium, namely Gomphostilbia (12 species), Nevermannia (5 species), Montisimulium (1 species), Simulium sensu stricto (21 species), and an unknown subgenus with one species (Simulium baimaii). The length of the ITS2 ranged from 247 to 308 bp. All black fly species had high AT content, ranging from 71 to 83.8%. Intraindividual variation (clonal variation) occurred in 13 species, ranging from 0.3 to 1.1%. Large intrapopulation and interpopulation heterogeneities exist in S. feuerboni from the same and different locations in Doi Inthanon National Park, northern Thailand. Phylogenetic relationships among 40 black fly species were examined using PAUP (version 4.0b10) and MrBAYS (version 3.0B4). The topology of the trees revealed two major monophyletic clades. The subgenus Simulium and Simulium baimaii were placed in the first monophyletic clade, whereas the subgenera Nevermannia + Montisimulium were placed as the sister group to the subgenus Gomphostilbia in the second monophyletic clade. Our results suggest that S. baimaii belongs to the malyschevi-group or variegatum-group in the subgenus Simulium. The molecular phylogeny generally agrees with existing morphology-based phylogenies.     

First report of a trichomycete fungus (Zygomycota: Trichomycetes) inhabiting larvae of Simulium ochraceum sensu lato Walker (Diptera: Simuliidae) from the Galapagos Islands

A collection of the larval black fly Simulium ochraceum sensu lato Walker was made at six lotic habitats on San Cristobal Island, Galapagos Islands, Ecuador, during May 2003. Sixty-eight larvae were assayed for the presence of gut fungi (Zygomycota: Trichomycetes). A trichomycete, designated as Smittium sp. was found inhabiting the hindgut of 17.6% of S. ochraceum s.l. larvae. Implications of this finding with respect to both host and trichomycete biogeography are discussed.     

Agricultural land use and black fly (Diptera,Simuliidae) species richness and species assemblages in tropical streams,Northeastern Thailand

Habitat degradation through agricultural land use is the major factor threatening lotic ecosystems. Although black flies are major components of these ecosystems, the impact of agricultural land use on species diversity and species assemblages has been largely ignored in tropical streams of the Oriental region. The objectives of this study are to examine patterns of species distribution and species richness and to compare black fly species richness and species assemblages in forest and agricultural streams in Thailand. A total of 143 collections were made from 70 stream sites between June 2007 and May 2008. Whereas 19 black fly species found in these collections were all found in forest sites, only 13 species were found in agricultural sites. High species richness was associated with larger, faster, and cooler streams with larger streambed particles and the presence of riparian trees. Logistic regression analyses revealed that stream size, velocity, and riparian vegetation are among the most important factors determining patterns of spatial distribution. The results are largely consistent with studies in other zoogeographic regions, suggesting the existence of general rules for black fly species distributions. Comparisons of the physicochemical conditions between forest and agricultural streams indicated that streams in agricultural areas are warmer, with higher conductivity and fewer riparian trees. Species richness was significantly higher in forest than in agricultural streams (t = 3.61, P < 0.001). Streams in forest areas were predominantly occupied by S. siamense (73%) but other species were also found at a relatively high frequency (>20%) of the sampling sites. In contrast, streams in agricultural areas were predominantly occupied by S. aureohirtum (>80%) among the sole black fly species at 27% of the sites. The results indicate that agricultural land use has a significantly detrimental impact on black fly diversity and species assemblages. Handling editor: D. Dudgeon     

Diversity, distribution and larval habitats of North Swedish blackflies (Diptera: Simuliidae)

1. Blackfly species richness and community structure were analysed at fifty-six sites in northern Sweden in two seasons. The sites were situated in a wide range of streams and rivers from small springbrooks, bog streams and lake-outlet streams to medium-sized forest rivers and large rivers draining montane regions.
2. Thirty-nine blackfly species were found, with between two and thirteen species per site. Neither species richness nor abundance could be related to the environmental variables measured.
3. An analysis of labral fan size of blackflies indicated a clear trend for the prevalence of larvae with small fans in large rivers and larvae with larger fan size in small streams. Similarly, fan size related to current velocities so that large fans were associated with slow current velocities and small fans with high velocities.
4. A strong relationship existed between species composition and habitat, as seen in ordination by non-metric multidimensional scaling. The relationship found between fan size and habitat size-related variables, such as channel width, depth, velocity and substratum particle size, along with longitude and altitude, in partial least squares regression analysis offered an explanation of the species composition–habitat relationship.
5. In addition to testing that distributions of blackfly larvae reflect morphological traits, we tested two general hypotheses pertaining to distribution patterns: (a) that blackfly communities show bimodal distributions; and (b) that their distributions are nested. Neither of these two hypotheses was supported by our observations. However, widespread blackfly species were locally more abundant than those found at relatively few sites, thus showing a positive abundance–occupancy relationship.     

Flow patterns around cocoons and pupae of black flies of the genusSimulium (Diptera : Simuliidae)

Flow patterns around structurally different cocoons and pupae of five species ofSimulium Latreille are described. Three features of the flow pattern common to all cocoons are; 1) a solenoidal vortex around the cocoon, 2) upward flow anterior (downstream) to the cocoon, and 3) one or two pairs of spiral-shaped vortices, which either touch or envelop the fill filaments of the pupa. The solenoidal vortex and the upward-spiralling, downstream vortices are common features of flow patterns around most bluff bodies submerged in a boundary layer. The proximity of the vortices to the fill filaments of all pupae suggests that these vortices are associated with gaseous exchange at the gill filaments.     

Variation in larval black fly populations at three sites in a stream system over five years (Diptera:Simuliidae)

Five years of monthly samples taken from three sites in the Broad Cove River, two at lake outlets and one a kilometer downstream from an outlet were analyzed. Considerable variations in the population of each species between sites and years were observed. Fluctuations in the populations were not always consistent from site to site or between species at the same site from one year to the next. As all sites were on the same stream and within 3 km of each other with only slight altitude differences climatic factors were not a major factor producing variations observed between sites. Also the previous years population level did not appear to be directly related to the following seasons population level. Apparently localized conditions at a site were exerting a marked influence on the larval simuliid population at each site.