Biological warfare in the garden pond: tadpoles suppress the growth of mosquito larvae

Abstract. 1. Although tadpoles and mosquito larvae may compete for scarce resources in natural freshwater systems, the mechanisms involved in such competition remain largely unstudied.
2. Replicated artificial ponds were set up to examine the role of pathogenic interference (water-borne growth inhibitors) in two tadpole–mosquito systems from south-eastern Australia. One system comprised taxa that are commonly sympatric in freshwater ponds (tadpoles of Limnodynastes peronii and larvae of Culex quinquefasciatus ) while the other comprised species that co-occur in brackish water ponds (tadpoles of Crinia signifera and larvae of Ochlerotatus australis ).
3. Water that had previously contained tadpoles suppressed the rates of survival and pupation of mosquito larvae in both systems. Fungicide reduced or eliminated this effect, suggesting that the growth inhibitors may be fungal organisms (possibly the yeast Rhodotorula glutinis ) from tadpole faeces. Fungicide also enhanced growth rates of tadpoles.
4. These results suggest that interference competition between tadpoles and mosquito larvae is mediated by other organisms in some ecological systems.     

Oviposition site selection by mosquitoes is affected by cues from conspecific larvae and anuran tadpoles

Abstract The oviposition site that a female mosquito selects will influence the fitness of her larvae. We conducted a series of artificial pond experiments to compare the oviposition responses of two species of mosquitoes with the presence of tadpoles, conspecifics and chemical cues from these organisms. The two mosquito species differ markedly in larval ecology. The larvae of one species, Culex quinquefasciatus, co‐occur with numerous freshwater organisms, including tadpoles of Linmodynastes peronii (the striped marsh frog). Larvae of the other mosquito, Ochlerotatus australis, inhabit small brackish rock ponds where the main potential competitors are tadpoles of Crinia signifera (the common eastern froglet). In field trials, females of both mosquito species oviposited significantly more often in water that contained (or had previously contained) conspecific larvae. However, these superficially similar responses were mediated via different pathways: fungicide abolished the response by C. quinquefasciatus but not by O. australis. The two mosquito species also responded differently to cues associated with syntopic tadpoles. The presence of tadpoles did not influence oviposition by C. quinquefasciatus, but O. australis oviposited less often if tadpoles were present. These interspecific differences in oviposition behaviour may be adaptive to differences in larval ecology: competition with tadpoles is likely to be more significant for O. australis than for C. quinquefasciatus. Our findings thus support the hypothesis that mosquitoes oviposit selectively to avoid potential anuran larval competitors.     

Competition between tadpoles and mosquito larvae

Tadpoles and mosquito larvae often co-occur, and may compete for scarce resources. However, competition between such distantly related organisms has attracted less scientific attention than have interactions among closely related taxa. We examined ecological interactions in two tadpole-mosquito systems in southeastern Australia, one from freshwater ponds (Limnodynastes peronii and Culex quinquefasciatus) and one from brackish-water habitats (Crinia signifera and Ochlerotatus australis). Diets of these tadpoles and mosquito larvae overlap considerably, potentially leading to competition for food. Laboratory experiments show that, in both study systems, mosquitoes reduced the growth rates of tadpoles, and tadpoles reduced the growth rates and survival of mosquito larvae. These negative effects were seen even at high food levels. Thus, our study suggests that tadpoles and mosquito larvae affect each other strongly, and do so via pathways other than simple consumptive competition. Because mosquitoes are important vectors for human diseases, the global decline in amphibian populations may have more impact on human health than has generally been anticipated.     

Influence of resource levels,organic compounds and laboratory colonization on interspecific competition between the Asian tiger mosquito Aedes albopictus (Stegomyia albopicta) and the southern house mosquito Culex quinquefasciatus

The mosquitoes Aedes albopictus (Stegomyia albopicta) (Skuse) and Culex quinquefasciatus (Say) (Diptera: Culicidae) are common inhabitants of tyres and other artificial containers, which constitute important peridomestic mosquito breeding habitats. We tested the hypotheses that interspecific resource competition between the larvae of these species is asymmetrical, that the concentration of chemicals associated with decomposing detritus affects the competitive outcomes of these species, and that wild and colonized strains of Cx. quinquefasciatus are affected differently by competition with Ae. albopictus. We conducted two laboratory competition experiments wherein we measured survivorship and estimated population growth (λ′) in both species under multiple mixed‐species densities. Under varying resource levels, competition was asymmetrical: Ae. albopictus caused competitive reductions or exclusions of Cx. quinquefasciatus under conditions of limited resources. In a second experiment, which used both wild and colonized strains of Cx. quinquefasciatus, organic chemical compounds associated with decomposing detritus did not affect the competitive outcome. The colonized strain of Cx. quinquefasciatus had greater survivorship and adult mass, and faster development times than the wild strain, but both strains were similarly affected by competition with Ae. albopictus. Competition between these species may have important consequences for vector population dynamics, especially in areas in which tyres and artificial containers constitute the majority of mosquito breeding habitats.     

Drinking rate regulation in some fresh-water mosquito larvae

ABSTRACT. Drinking rates were determined with four species of freshwater mosquito larvae by colorimetric measurement of the dye ingested after groups of fourth instars were allowed access for set periods to 2% amaranth solutions. The rate of drinking for the saline-tolerant Aedes aegypti, 309±113 nl per h per individual, was comparable with rates given in the literature for several saline-water species, but rates for Culex quinquefasciatus, Culex molestus and Anopheles albimanus were markedly lower (167±30, 48±17 and 108±28m per h, respectively). When larvae of A.aegypti and C. quinquefasciatus were glutted with kaolin (allowed to replace all food in the gut by filter-feeding in kaolin suspension), drinking rates were little affected at first, but after 1 day of fasting (holding in water after glutting), drinking rates were 50% lower and were reduced by a further 20% with fasting for up to 3 days. For A.aegypti, C.quinquefasciatus and C. molestus, drinking rates were approximately doubled with kaolin dispersed in the dye solution, and after fasting, were increased by up to 100% in solutions containing 0.05% of water-soluble yeast extract. A similar phagostimulant effect of 10^-3M adenylic acid was demonstrated for A. aegypti and C. quinquefasciatus. A single experiment indicated similar stimulatory effects of kaolin and adenylic acid for, A. albimanus. With 0.01-0.05% agarose in the dye solutions, drinking rates for A. aegypti and C. quinquefasciatus were more than doubled, and a similar though weaker effect was demonstrated for another colloid, methylcellulose. In constrast, both colloids markedly reduced the rate of drinking with A.albimanus. These findings are discussed in relation to whether drinking and filter-feeding are necessarily coupled. The possible significance of this with respect to larvae that feed in different microhabitats, providing different levels of dissolved and colloidal nutrient organic matter, is considered. The implications of drinking rates for biotests of solubilized bacterial toxins as mosquito larvicides are noted.     

Using competitive and facilitative interactions in intercropping systems enhances crop productivity and nutrient-use efficiency

This paper reviews recent research on the processes involved in the yield advantage in wheat (Triticum aestivum L.)/maize (Zea mays L.), wheat/soybean [Glycine max (L.) Merr.], faba bean (Vicia faba L.)/maize, peanut (Arachis hypogaea L.)/maize and water convolvulus (Ipomoea aquatica Forsk.)/maize intercropping. In wheat/maize and wheat/soybean intercropping systems, a significant yield increase of intercropped wheat over sole wheat was observed, which resulted from positive effects of the border row and inner rows of intercropped wheat. The border row effect was due to interspecific competition for nutrients as wheat had a higher competitive ability than either maize or soybean had. There was also compensatory growth, or a recovery process, of subordinate species such as maize and soybean, offsetting the impairment of early growth of the subordinate species. Finally, both dominant and subordinate species in intercropping obtain higher yields than that in corresponding sole wheat, maize or soybean. We summarized these processes as the `competition-recovery production principle'. We observed interspecific facilitation, where maize improves iron nutrition in intercropped peanut, faba bean enhances nitrogen and phosphorus uptake by intercropped maize, and chickpea facilitates P uptake by associated wheat from phytate-P. Furthermore, intercropping reduced the nitrate content in the soil profile as intercropping uses soil nutrients more efficiently than sole cropping.     

The community ecology of Aedes egg hatching: implications for a mosquito invasion

Abstract. 1. A recently introduced treehole mosquito from Asia, Aedes albopictus , is spreading throughout eastern North America, especially in tyre-refuse piles. Previous studies have identified inhibitory effects of larvae on egg hatch as a potential population regulatory mechanism within Aedes. Larva-egg interactions may also occur between species. This experiment assesses the ability of larvae of A. albopictus and two possible competitors in North America, A.triseriatus and A.aegypti , to suppress hatching of conspecific and congeneric eggs.
2. We exposed eggs of each species to varying combinations of larval species and density for 24h and assessed subsequent hatch rates. Aedes albopictus eggs exhibited the lowest level of inhibition when exposed to high larval densities; moreover, at the lowest larval density they imposed the most intense interspecific hatch inhibition.
3. Discretionary hatching in response to larval density may influence community composition by promoting the spread of A.albopictus , perhaps even leading to its dominance within North American Aedes communities.     

Septic tank mosquitoes: competition between species in central Nigeria

Exit traps, placed over the air vents of septic tanks, were used to examine species diversity and relative abundance of mosquitoes breeding in ammonia-rich waters of septic tanks. Of the six species found, Culex decens Theobald and Culex cinereus Theobald appeared to be competing successfully with Culex quinquefasciatus Say during the wet season but not during the long dry season. The seasonal timing of their displacement by Cx quinquefasciatus was variable and did not correlate well with climatic factors. The three other species present, generally during the wet season and early dry season, were Culex tigripes G. & C., Culex horridus Edwards and Aedes aegypt (L.). Experimental bucket ovitraps were used to assess preference towards covered (dark) septic tank water in comparison with sunlit septic tank water, covered and sunlit compost water. These were colonized by Cx quinquefasciatus, Cx decens, Ae. aegypti and Ae. vittatus Bigot. The covered septic tank water was more abundantly colonized by Cx quinquefasciatus and marginally so by the two Aedes species. Cx decens appeared to colonize the exposed compost water more readily in the dry season, but changed to the covered septic tank water in the wet season. The discussion centres around competition between these mosquitos species and concludes that it would be useful to know what environmental factors, or what aspects of competition, lead to severe natural reductions in the abundance of the major pest species Cx quinquefasciatus.     

Temporal and spatial habitat preferences and biotic interactions between mosquito larvae and antagonistic crustaceans in the field

Investigations on natural antagonists of mosquito larvae found that micro‐crustaceans (e.g., Cladocera) control mosquito populations under experimental conditions. However, their relevance for mosquito control under field situations remains widely unclear because important information about habitat preferences and time of occurrence of crustaceans and mosquito larvae are still missing. In order to fill this knowledge gap, a field study was undertaken in different wetland areas of Saxony, Germany, in different habitats (i.e., grassland, forest, and reed‐covered wetlands). We found negative interactions between larvae of Ae. vexans and predatory Cyclopoida (Crustacean: Copepoda), which both were dominant during the first two weeks of hydroperiod, at ponds located at grassland habitats. Larvae of Cx. pipiens were spatially associated with competing Cladocera, but they colonized ponds more rapidly. Populations of Cladocera established from the third week of hydroperiod and prevented Cx. pipiens colonization thereafter. Ostracoda were highly abundant during the whole hydroperiod, but their presence was restricted to habitats of reed‐covered wetland at one geographical area. Mosquito larvae hardly occurred at those ponds. In general, we found that ponds at the reed‐covered wetlands provided better conditions for the initial development of crustaceans and hence, mosquito larval colonization was strongly inhibited. Grassland habitat, in contrast, favored early development of mosquito larvae. This study showed that micro‐crustaceans are relevant for mosquito management but their impact on mosquito larvae varies between species and depends on environmental conditions.     

Priority effects of bamboo-stump mosquito larvae: influences of water exchange and leaf litter input

Abstract 1. A series of laboratory experiments was conducted to examine the influences of leaf litter input and flushing of medium on the priority effects of the two bamboo-stump breeding mosquitoes, Aedes albopictus and Tripteroides bambusa , using 150-ml microcosms. Larvae of either species were introduced to microcosms on day 0 (early cohort), at different densities, and day 14 (late cohort). The effects of the early cohort on survival, pupation, and biomass yield of the late cohort were compared among various combinations of the two species and among different treatments (water exchange, leaf addition, both of them, neither of them) on days 14 and 44.
2. Survivorship and pupation success of the late A. albopictus cohort were affected negatively by the presence of either species of the early cohort to a greater extent than those of the late T. bambusa cohort.
3. Water exchange reduced mortality of the late A. albopictus cohort over a short term in the presence of the early cohort of either species, indicating that a toxic substance was involved in the inhibitory priority effects. The addition of leaf litter enhanced survivorship and pupation of the late A. albopictus cohort, whether or not water was exchanged. The late T. bambusa cohort showed high survival rate with all treatments.
4. The results indicate that leaf-litter input moderates the inhibitory priority effects on A. albopictus larvae, not only by supplying food resources but also by alleviating the toxicity of accumulated dissolved substances.     

When carnivores collide: a review of studies exploring the competitive interactions between bobcats Lynx rufus and coyotes Canis latrans

1. Bobcats Lynx rufus and coyotes Canis latrans are two widespread mesopredators with a complex history of sympatry. The competitive interactions between these species are of interest to biologists due to the furbearer status of bobcats, the recent range expansion of coyotes, and the recolonisation of several parts of North America by bobcats following their extirpation. Although studies exploring the dynamics and competition between bobcats and coyotes span decades, there is a lack of understanding regarding what factors influence exploitative or interference competition, and what methodologies are conducive to identifying these types of competition.
2. We gathered a comprehensive list of research papers (n = 41) exploring bobcat–coyote competitive interactions in North America. From them, we collected the following: study site characteristics, number and types of research methods, number and types of metrics explored, history of sympatry of the two species at the study location, presence of apex predators, and documentation of interference and/or exploitative competition.
3. Using generalised linear models, we determined that interference competition between bobcats and coyotes was observed primarily in open habitat. However, habitat heterogeneity, the number of research methods and metrics used, presence of an apex predator, and history of sympatry could not be used to predict the occurrence of interference competition. Studies that included diet overlap were less likely to observe interference competition than studies that used other metrics to infer competition.
4. Competitive interactions between coyotes and bobcats are largely a function of prey availability. Our findings suggest that habitat type may be a surrogate for prey availability, which many researchers mention, but do not explicitly measure. Future studies investigating bobcat–coyote interactions should include the quantification of prey densities to gain a more comprehensive understanding of the system at large, and should avoid using solely diet or habitat overlap as metrics to assess competition.

     

Bamboo stumps as mosquito larval habitats in Darjeeling Himalayas, India: A spatial scale analysis

Bamboo stumps can be a congenial breeding habitat of the mosquitoes. In view of this, a preliminary assessment of the dipteran immatures inhabiting the stumps of bamboo groves in the Darjeeling Himalayas was carried out at a spatial scale. Of the 104 stumps of Dendrocalamus hamiltoni surveyed, 70 were found to host immatures of three dipteran species, the mosquitoes Aedes aegypti and Culex quinquefasciatus and the midges Chironomus sp. in varying densities. Though the stumps varied in diameter, in each stump on average 12. 1 immatures were found. The abundance of the immatures was positively correlated with the diameter of the stumps (r=+0.382; P < 0.001) but negatively with the pH of the water present in the stumps (r=–0.336; P < 0.01). The coefficient of association was found to be +8.4 for the Ae. aegypti and Chironomus immatures, while in the rest of the species pair the association seemed to be independent. Thus it can be concluded that the stumps in the bamboo groves of Darjeeling Himalayas provides a favourable habitat for the mosquito and chironomid immatures.     

The influence of ecological factors on mosquito abundance and occurrence in Galápagos

We sampled mosquitoes across 18 sites established at different elevations and stretching from the north to the south of Isla Santa Cruz, Galápagos. Two commonly occurring species, Ae. taeniorhynchus and Cx. quinquefasciatus, were collected along with environmental variables characteristic of the trapping sites to assess their influence on mosquito abundance and occurrence in the dry season of 2015. We captured Ae. taeniorhynchus at 14 out of 18 sites and Cx. quinquefasciatus at low and high elevation sites on Santa Cruz. We utilized two generalized linear models; the first assessed the influence of environmental variables on abundances of Ae. taeniorhynchus and the second assessed the influence of these variables on the presence of Cx. quinquefasciatus. Populations of both mosquito species declined with elevation. Rainfall data were limited, as we sampled during the dry season of 2015. Distance to mangroves and maximum humidity were significant in influencing the abundance of Ae. taeniorhynchus, while maximum humidity was found to significantly influence the presence of Cx. quinquefasciatus. Both species occurred in sites where temperature, precipitation, and humidity should allow for mosquito development as well as parasitic development of the protozoan parasites that cause avian malaria. Further research involving year‐round sampling of mosquitoes and accompanying meteorological data as well as experimental studies on vector competence are required to understand disease dynamics of parasites such as avian malaria in Galápagos.     

The comparative activity of the Australian and united states strains of Culicinomyces clavosporus bioassayed in mosquito larvae of three different genera

Anopheles hilli, Culex quinquefasciatus, and Aedes aegypti were used as test insects to compare the activity of the Australian and United States strains of Culicinomyces clavosporus. To minimize the variability incurred by using different larval batches, both strains were bioassayed at the same time using one batch of larvae. Six pairs of assays for each of the three test species were conducted in this manner. It was found that there was no difference in potency of the two strains in any one of the three species. A between species comparison, with the data pooled for both strains, showed that A. aegypti was more susceptible to the fungus than A. hilli. The susceptibility of C. Quinquefasciatus appeared to be intermediate but the fiducial limits of the weighted mean LC₅₀ overlapped with those of the other two species. From the results of these experiments it would seem that, with regard to potency, both strains of Culicinomyces may be equally promising for the biological control of mosquitoes.     

Satyrization and satyrization‐resistance in competitive displacements of invasive mosquito species

Competitive displacements or reductions of resident populations of insects, often effected by a related species, may be caused by a variety of mechanisms. Satyrization is a form of mating interference in which males of one species mate with females of another species, significantly decreasing their fitness and not generating hybrids. Satyrization has been established to be the probable cause of competitive displacements of resident mosquitoes by invasive species, especially of Aedes aegypti by Aedes albopictus, two important vectors of dengue and chikungunya viruses. Mathematical models predict that even low levels of asymmetric mating interference are capable of producing competitive displacements or reductions. Couplings of virgin Ae. aegypti females with Ae. albopictus males effectively sterilize these females through the monogamizing actions of male accessory gland products, but the converse interspecific mating does not impact the future reproduction of Ae. albopictus females. Populations of Ae. aegypti exposed to satyrization quickly evolve resistance to interspecific mating, which is believed to ameliorate reproductive interference from, and promote co‐existence with, Ae. albopictus. The evolution of satyrization resistance among Ae. aegypti in laboratory cages is accompanied by fitness costs, such as reduced fecundity and slower receptivity to conspecific males. Cage experiments and field observations indicate that Ae. albopictus males are capable of satyrizing females of other species of the Stegomyia subgenus, potentially leading to competitive displacements, and possible extinctions, especially of endemic species on islands. Examination of other examples of reproductive interference in insects reveals few parallels to the mechanism and outcomes of satyrization by Ae. albopictus. We conclude by posing the hypothesis that satyrization may favor the ecological success of Ae. albopictus, and suggest many lines for future research on this phenomenon.     

Oviposition fluids from adult wasps mediate interspecific competition between parasitoid larvae

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Effects of temperature and diet on development and interspecies competition in Aedes aegypti and Aedes albopictus

We asked whether climate change might affect the geographic distributions of Aedes aegypti (L.) and Aedes albopictus (Skuse) (Diptera: Culicidae). We tested the effects of temperature, diet and the presence of congeneric species on the performance of immature stages of these two aedine species in the laboratory. Mosquitoes in three different species-density combinations were reared at four constant temperatures (20 °C, 25 °C, 30 °C, 35 °C) on low- or high-level diets. Of the four temperatures tested, mortality increased only at 35 °C in both species. Mortality was higher on the high-level diet than on the low-level diet at 35 °C, but not at other temperatures. The presence of congeneric species had a significant positive effect on mortality in Ae. albopictus, but not in Ae. aegypti. Both species developed more quickly at higher temperatures within the range of 20-30 °C; development was not enhanced at 35 °C. Population growth of Ae. albopictus was more stable, regardless of diet and temperature; that of Ae. aegypti varied more according to these two factors. These species-specific attributes may help to explain the latitudinal distribution of the mosquitoes and degree of species dominance where they are sympatric.     

The naturally derived insecticide spinosad is highly toxic to Aedes and Anopheles mosquito larvae

Spinosad is a naturally derived biorational insecticide with an environmentally favourable toxicity profile, so we investigated its potency against mosquito larvae (Diptera: Culicidae). By laboratory bioassays of a suspension concentrate formulation of spinosad (Tracer), the 24 h lethal concentration (LC50) against Aedes aegypti (L.) third and fourth instars was estimated at 0.025 p.p.m. following logit regression. The concentration-mortality response of third- and fourth-instar Anopheles albimanus Weidemann did not conform to a logit model. The LC50 value of spinosad in Anopheles albimanus was 0.024 p.p.m. by quadratic linear regression. A field trial in southern Mexico demonstrated that spinosad 1 p.p.m. compared with the standard temephos (Abate) 1% granules 100 g/m3 water prevented Ae. aegypti breeding in plastic containers of water for 8 weeks; at 10 p.p.m. spinosad prevented breeding for > 22 weeks. In another field trial, spinosad at 5 p.p.m. and temephos both completely eliminated reproduction of Ae. aegypti for 13 weeks. In contrast, the bacterial insecticide Bacillus thuringiensis var. israelensis (Bti, Vectobac) AS) performed poorly with just 2 weeks of complete inhibition of Ae. aegypti breeding. Spinosad also effectively prevented breeding of Culex mosquitoes and chironomids in both trials to a degree similar to that of temephos. We conclude that spinosad merits evaluation as a replacement for organophosphate or Bti treatment of domestic water tanks in Mesoamerica. We also predict that spinosad is likely to be an effective larvicide for treatment of mosquito breeding sites.     

Synergism between permethrin and propoxur against Culex quinquefasciatus mosquito larvae

To see if synergism occurs between carbamate and pyrethroid insecticides, we tested permethrin and propoxur as representatives of these two classes of compounds used for mosquito control. Larvicidal activity of both insecticides was assessed separately and together on a susceptible strain of the mosquito Culex quinquefasciatus (Diptera: Culicidae) by two methods. When mixed at a constant ratio (permethrin : propoxur 1 : 60 based on LC50) and tested at serial concentrations to plot dose/mortality regression, significant synergy occurred between them (co-toxicity coefficient = 2.2), not just an additive effect. For example, when the mixture gave 50% mortality, the same concentrations of permethrin and propoxur alone would have given merely 2 x 1% mortality. When a sublethal dose (LC0) of permethrin or propoxur was added to the other (range LC10-LC95), synergism occurred up to the LC80 level. Synergistic effects were attributed to the complementary modes of action by these two insecticide classes acting on different components of nerve impulse transmission. Apart from raising new possibilities for Culex control, it seems appropriate to consider using such mixtures or combinations for insecticide-treated mosquito nets in situations with insecticide-resistant Anopheles malaria vectors.