Seasonal variation of black fly (Diptera: Simuliidae) species diversity and community structure in tropical streams of Thailand

Seasonal variation of the physicochemical conditions of streams plays an important role in aquatic insect diversity and community structure. Asian tropical streams profoundly change between seasons due to the effects of monsoons. However, little is known about how these changes affect aquatic insect diversity and community structure. The objectives of this study are to examine seasonal variations of the physicochemical conditions in tropical streams in Thailand and to assess the effects of these changes on black fly community structure and diversity. Black flies were sampled and physicochemical conditions recorded at eight sites between December 2007 and December 2008. A total of 10 black fly species were found. Comparisons of the streams between seasons revealed that physical conditions related to rainfall rate were significantly different. Canonical correspondence analysis differentiated sampling sites from each season. Streams in the rainy season were faster and deeper, with higher discharge and conductivity than those of the cold and dry seasons. Species richness was significantly higher in the rainy season than in the cold and dry seasons ( F  = 6.23, P  = 0.004). Community structure profoundly changed between the low-flow season (cold and dry) and high-flow season. Black fly species found predominantly in the low-flow season ( Simulium siamense "cytoform A", S. aureohirtum ) decreased dramatically during the high-flow season. In contrast, species found at high frequency during the high-flow season ( S. nakhonense , S. angulistylum ) disappeared in the low-flow season. The study demonstrates the importance of seasonal variation of stream conditions on black fly community structure and diversity.     

Black fly larvae facilitate community recovery in a mountain stream

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Disturbance to a stream food web by a bacterial larvicide specific to black flies: feeding responses of predatory macroinvertebrates

1. A field study was conducted to: (i) assess feeding habit changes of two predatory stoneflies following the loss of larval black fly (Diptera: Simuliidae) prey from two streams; and (ii) determine the relative importance of black fly larvae as prey for these and other selected predatory benthic macroinvertebrates. 2. Acroneuria lycorias and Paragnetina media (Plecoptera: Perlidae) diets were monitored in response to local reductions in larval black fly populations caused by Bacillus thuringiensis var. israelensis (B.t.i.) in two Michigan streams. These predators were collected from B.t.i.-treated and control sections of the streams, and their foreguts inspected for prey. 3. Black flies were the major dietary component of both predators collected from the control sections, but the number of black flies ingested was significantly less for predators collected from B.t.i.-treated habitats. Total number of prey ingested significantly decreased for A. lycorias, but not for P. media, and non-black fly prey consumption significantly increased for P. media, but not for A, lycorias, following B.t.i. applications. 4. In prey choice trials conducted in experimental channels, A. lycorias and P. media showed no preference between prey types (black flies and mayflies). Body mass gain of individual A. lycorias nymphs was measured, and was similar for nymphs in black fly-rich and black fly-poor environments. Conversely, Isoperla signata and I. dicala (Plecoptera: Perlodidae) ingested significantly more Simulium vittatum (Diptera: Simuliidae) than Baetis flavistriga (Ephemeroptera: Baetidae) or Epeorus vitrea (Heptageniidae) prey. Boyeria vinosa (Odonata: Aeshnidae) ingested significantly more B. flavistriga than S. vittatum prey. 5. Reducing black fly densities in these streams, using B.t.i., indirectly and differentially affected predators. In black fly-poor environments, feeding habits of specialist predators were most affected, and generalist predators least affected because the latter consumed alternative prey. Predator—predator and predator-prey interactions, and prey community structure may be affected indirectly by disturbances such as B.t.i. applications by reducing food resources and forcing predation onto less preferred prey.     

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

A new species of microsporidium, Caudospora palustris (Microsporidia: Caudosporidae), is described from 3 species of black flies (Cnephia ornithophilia and diploid and triploid cytospecies of Stegopterna mutata), bringing to 7 the total species of caudosporids recorded from North America. This new species of caudosporid is recorded from swamp streams of the Coastal Plain from New Jersey to Georgia, with single records from the New Jersey mountains and the Upper Peninsula of Michigan. Densities of patently infected larvae (up to 10,600/m2) and spore production (nearly 8x10(11)/m2) are the greatest recorded for any microsporidium of black flies. The ultrastructure of this new species is presented, along with the first molecular characterization for a microsporidium of black flies. The phylogenetic position of black fly microsporidia within the phylum Microsporidia is presented; however, the analysis does not support the inclusion of C. palustris in any clade. Key features of all North American caudosporids are provided, and possible evolutionary trajectories are proposed based on optimization of caudosporid species on the phylogeny of their 22 known host species, including 16 that represent new host species records.     

Alpha richness and niche breadth of a widespread group of aquatic insects in Nearctic and Neotropical streams

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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.     

The roles of abiotic factors, dispersal, and species interactions in structuring stream assemblages of black flies (Diptera: Simuliidae)

ABSTRACT: BACKGROUND: The patterns and drivers of species assemblages represent the core of community ecology.We focus on the assemblages of a single family of ubiquitous lotic insects, the Simuliidae(black flies), of which the larvae play a critical role in resource turnover in steams. We useMantel tests and null models to tease out the potential influence of abiotic stream conditions,species interactions, and dispersal on the assemblage patterns of larval black flies over twospatial scales (within and across ecoregions) and two seasons (spring and summer). RESULTS: When stream sites were considered across ecoregions in the spring, stream conditions anddispersal were correlated significantly with species similarity; however, within ecoregions inthe spring, dispersal was important only in the Piedmont and Sandhills and abiotic factorsonly in the Mountains. In contrast, results of the summer analyses within and acrossecoregions were congruent; assemblage similarity was significantly correlated with streamconditions both across and within ecoregions. Null models suggested that patterns of speciessegregation in the spring were consistent with a community structured by competition,whereas patterns in the summer were consistent with species assemblages influenced byabiotic factors. CONCLUSIONS: Species composition of black flies at streams sites is correlated with dispersal factors andstream conditions, but results vary over spatial and temporal scales. Communities of blackflies can be viewed within a metacommunity context; local assemblages are consistent withspecies sorting and mass effects. Given that black flies have a terrestrial stage, with femalesdeciding where to place the eggs, a full understanding of the processes that determine local aquatic assemblages will require integration of the dynamics of the aquatic immature stagesand the terrestrial adults.     

Seasonality of trichomycetes in larval black flies from South Carolina, USA

Trichomycete fungi are common endobionts of aquatic insect larvae, but little is known of their ecology. In this study, the seasonality of trichomycete colonization of larval black flies (Diptera: Simuliidae) was investigated in three streams in northwestern South Carolina. At least eight species of trichomycetes were found in two species of black flies, and 93.8% of 1819 larval black flies examined contained trichomycetes. Significant differences were found in the seasonal prevalence of Harpella melusinae, Simuliomyces microsporus, and Paramoebidium spp. at one of three sites. At this site, the lowest mean prevalence for H. melusinae occurred in winter (67%) versus the other seasons (96-100%), whereas mean prevalence was lowest in summer for Simuliomyces microsporus (1%) versus the other seasons (2-21%) and lowest in summer for P. spp. (9%) versus the other seasons (45-67%). Significant differences in levels of colonization among seasons were not detected. Conjugations of H. melusinae (representing early stages of sexual reproduction) occurred most frequently in the spring and winter (up to 14% of larvae). Sexual reproduction (represented by zygospores) of Legeriomycetaceae occurred most frequently in the spring and fall (up to 17% of larvae).     

Fly pollination of Gorteria diffusa (Asteraceae), and a possible mimetic function for dark spots on the capitulum

We investigated the functional significance of raised black spots on the ray florets of Gorteria diffusa (Asteraceae) in South Africa. Field observations showed that G. diffusa is pollinated by a small bee-fly, Megapalpus nitidus (Bombyliidae), which is strikingly similar to the raised spots that occur on some of the ray florets. Removal of the spots resulted in a significant decrease in the rate of fly visits to capitula, but did not significantly affect seed set. Replacement of the spots with simple ink spots also significantly reduced the rate of pollinator visits, suggesting that flies respond to details in the structure of the spots. Investigations using scanning electron microscopy showed that the spots of G. diffusa consist of a complex of different cell types. Differences in epidermal sculpturing may partly explain the UV reflectance pattern of these spots, which is similar to that of the flies. Male flies are strongly attracted to the spots, as well as to other flies sitting in the capitula, although female flies also visit the capitula. We conclude that the spots of G. diffusa mimic resting flies, thereby eliciting mate-seeking and aggregation responses in fly pollinators. Similar dark spots have evolved in unrelated South African Gazania, Dimorphotheca, and Pelargonium species pollinated by bee-flies.     

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     

Zoogeographic analysis of the fauna of the family Asilidae (Diptera) of the Lower Volga area

Based on the author’s own and the literature data, the zoogeographic analysis of robber flies (Asilidae) of the Lower Volga area was carried out for the first time. A total of 21 range types were distinguished and the fractions of different chorological groups in the fauna were determined. The zoogeographic structure of faunal complexes of robber flies inhabiting biotopes with different plant associations was studied.     

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.     

On the role of blue shadows in the visual behaviour of tsetse flies

Tsetse flies (Glossina spp.), the vector for African trypanosomiasis, are highly attracted by blue and black surfaces. This phototactic behaviour has long been exploited to trap tsetse flies as one measure in the control of African trypanosomiasis. However, why blue and black are so attractive for tsetse flies is still unknown. We propose that the combination of blue and black is attractive for many Glossina species because when searching for a shady resting place to pass the day, the flies are probably guided by the blueness and darkness of daytime shadows. In contrast to people's experience that daytime shadows are colourless, actually on a sunny day all shadows are tinted bluish by the scattered blue skylight.     

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.     

Distribution of amphibiotic insects of different trophic groups in mountainous and steppe rivers of Western Tuva

The distribution of preimaginal stages of amphibiotic insects (mayflies, stoneflies, caddisflies, and black flies) in the running waters of the mountain-steppe landscapes of western Tuva has been investigated. The basin of the Hemchik River is 500–2200 meters above sea level. The taxonomic composition and spatial distribution are determined; higlnnountain, middle- mountain and low-nountain plain types of communities are detailed. The trophic structure of amphibiotic communities is analyzed along the ecological profiles from the upper to the lower reaches of the rivers.     

The spatial distribution of black flies (Diptera: Simuliidae) in the basin of the Sema mountain river in the North Altai mountain region

The distribution of preimaginal phases of black flies in the basin of the mountain river Sema (North Altai) is analyzed based on quantitative data. The studies were carried out at the altitudes of 300–2500 above sea level. The structural elements of the black fly community in the altitudinal belts were distinguished. The distribution of black flies on the landscape and ecological profiles is discussed. The sequence of changes in the black fly community along the river is considered. An assessment of the importance of the environmental factors determining the unevenness of the black fly population in the Sema River basin is given.     

Fine structure of frozen-etched lipid granules in the fat body of an insect

Lipid storage in fat-body cells of adult female black flies was examined using freeze-etching electron microscopy. Frozen-etched lipid granules exhibited a laminated structure. The molecular arrangement of the lipid granule may depend on the physiological condition of the insect and may be involved in the control of lipid metabolism in the fat-body cell.     

Effects of eucalyptus afforestation on leaf litter dynamics and macroinvertebrate community structure of streams in Central Portugal

To test the hypothesis whether afforestation with Eucalyptus globulus affects litter dynamics in streams and the structure of macroinvertebrate aquatic communities, we compared streams flowing through eucalyptus and deciduous forests, paying attention to: (i) litterfall dynamics, (ii) accumulation of organic matter, (iii) processing rates of two dominant leaf species: eucalyptus and chestnut, and (iv) macroinvertebrate community structure. The amount of allochthonous inputs was similar in both vegetation types, but the seasonality of litter inputs differed between eucalyptus and natural deciduous forests. Eucalyptus forest streams accumulated more organic matter than deciduous forest streams. Decomposition of both eucalyptus and chestnut leaf litter was higher in streams flowing through deciduous forests. The eucalyptus forest soils were highly hydrophobic resulting in strong seasonal fluctuations in discharge. In autumn the communities of benthic macroinvertebrates of the two stream types were significantly different. Deciduous forest streams contained higher numbers of invertebrates and more taxa than eucalyptus forest streams. Mixed forest streams (streams flowing through eucalyptus forests but bordered by deciduous vegetation) were intermediate between the two other vegetation types in all studied characteristics (accumulation of benthic organic matter, density and diversity of aquatic invertebrates). These results suggest that monocultures of eucalyptus affect low order stream communities. However, the impact may be attenuated if riparian corridors of original vegetation are kept in plantation forestry.