Negative effects of habitat drying and prior exploitation on the detritus resource in an ephemeral aquatic habitat

Ephemeral aquatic habitats are characterized by cycles of drying and subsequent inundation, and by production of sequential non-overlapping cohorts of organisms. Both processes may alter the quantity or quality of resources, and may therefore affect survival and development of cohorts that subsequently colonize ephemeral habitats. We examined these effects of habitat drying and non-overlapping cohorts on experimental cohorts of the tree hole mosquito Aedes triseriatus, testing specifically whether the value of leaf litter as a food resource is altered by cycles of inundation and drying, or by exploitation by a prior non-overlapping cohort. We created four treatments of leaf litter: (1) no prior cohort, continuously wet; (2) no prior cohort, one␣wet/dry cycle; (3) prior cohort, continuously wet, and (4) prior cohort, one wet/dry cycle, and tested for effects on individual fitness components (survivorship, mean dry mass at, and median days to eclosion) and on population growth (estimated finite rate of increase –λ′). Both resource drying and the presence of a prior cohort negatively affected individual fitness components in tires, increasing days to eclosion, and decreasing mean dry mass at eclosion for both sexes. Resource drying also negatively affected estimated rates of increase (λ′) in tree holes. A prior cohort had no significant effects on λ′. These results indicate that intraspecific interactions among mosquito larvae may include amensalistic effects of earlier, non-overlapping cohorts, and that resource drying reduces resource quality. The latter effect indicates that enhanced production of A. triseriatus from recently filled containers is not due to resource drying per se, and may result from more complex community-level effects of habitat drying. Extreme cycles of drying and inundation seem likely to increase intraspecific resource competition among drought-adapted species like A. triseriatus. Received: 5 May 1997 / Accepted: 20 January 1998     

Competition and resistance to starvation in larvae of container-inhabiting Aedes mosquitoes

Abstract. 1. Hypotheses about declining populations of container-inhabiting Aedes mosquitoes following the invasion by additional species were tested.
2. The larval competition hypothesis was studied experimentally in pure and mixed cultures of Aedes aegypti (L.), A.albopictus (Skuse) and A.triseriatus (Say). The experiments used decomposing leaf litter in the laboratory, as opposed to most previous research which used non-natural food.
3. Resistance to starvation is introduced as a new measure of larval performance and competitiveness. The hypothesis is that more successful larvae store larger energy reserves and resist the lack of food longer.
4. Contrary to previous research showing better performance of A.aegypti in mixed cultures, A.albopictus developed faster and had greater survival when natural food was used.
5. Resistance to starvation was greater in the better performing species (i.e. A.aegypti with non-natural food and A.albopictus with leaf litter). Oxygen consumption by starved larvae was similar in the three container species, and in the ground-water mosquito, A.taeniorhynchus (Wied.), whose resistance to starvation was comparatively very low.     

Midgut and salivary gland barriers to La Crosse virus dissemination in mosquitoes of the Aedes triseriatus group

Vector competence for La Crosse virus (LACV) was compared for four species in the Aedes (Protomacleaya) triseriatus group: Ae. triseriatus (Say), Ae. hendersoni Cockerell, Ae. zoosophus Dyar and Knab and Ae. brelandi Zavortink (Diptera: Culicidae). Rates of replication and dissemination of virus in the mosquito hosts were compared and rates of oral transmission of virus to suckling mice were determined. Barriers to virus dissemination which limited the ability of each species to transmit virus were identified. Ae. zoosophus displayed the highest vector competence for LACV. Both infection and transmission rates were high: 99% and 85% respectively; no significant barriers to LACV were found. Disseminated infection of Ae. triseriatus with LACV was controlled primarily be a midgut escape barrier. When virus was introduced directly into the haemocoel, transmission rates were significantly increased (37% v. 79%). Ae. hendersoni showed high susceptibility to LACV infection but a very low rate of oral transmission (7%). Ae. brelandi was also highly susceptible to infection by LACV and transmitted virus at an intermediate rate (27%). Modulation of vector competence in both Ae. hendersoni and Ae. brelandi resulted from a salivary gland escape barrier. As these four species of mosquitoes comprise a closely related monophyletic series, their differences of vector competence for LACV provide an excellent model for studying the genetic basis of the barriers involved.     

Effects of crowding on populations of Aedes albifasciatus larvae under laboratory conditions

Aedes (Ochlerotatus) albifasciatus(Macquart 1838) (Diptera: Culicidae) is a neotropical flood water mosquito, incriminated as the main vector of the western equine encephalitis virus, and which affects beef and milk production in central Argentina. The short time required to hatch and develop from egg to adult, usually in temporary pools, suggests a strategy which allows for exploitation of transient pools, thus evading predation and interspecific competition. Under these conditions intra specific competition could represent a major density-dependent source of larval mortality, but the relative importance of density-dependent regulation of mosquito populations has generated controversy. Therefore we examined the effects of larval density on basic population characteristics of Ae. albifasciatusin the laboratory. Larvae were obtained by synchronous hatching of eggs laid by field-trapped females. Emerging larvae (L1) were used to build cohorts of different initial densities, kept in plastic trays with 400 ml of distilled water, and food supplied daily during the first 10 days (0.1 g per larvae day^–1). Age-specific development time and specific and relative mortality were estimated, and their relation to initial larval density was assessed through linear and non-linear regressions and correlation analysis. First hatching was registered 3 h after flooding the eggs. Higher levels of pre-adult mortality were detected in groups with higher densities. Specific mortality and average time to enter a stage of L1 to L3 could directly be related to initial larval density, but no significant relations were found for L4 and pupae. Results suggest that crowding could be a factor capable of regulating the density of natural populations of Ae. albifasciatus.     

Egg hatching inhibition: field evidence for population regulation in a treehole mosquito

ABSTRACT. 1. We investigated the influence of larval density on egg hatching in the treehole mosquito, Aedes triseriatus Say (Diptera: Culicidae) by submerging laboratory-produced egg batches in natural habitats stocked with fixed densities of larvae.
2. We found a significant decrease in hatching rate as larval density increased. The results suggest that inhibited eggs refrain from hatching into environments in which competitive effects of large larvae are potentially severe, and in which cannibalism may occur.
3. The amount of time the egg batches were exposed to experimental conditions (between 2 and 16 days) had no significant influence on hatch rate.     

Observations on wolbachiae in mosquitoes

A slightly modified Giemsa smear procedure consistently demonstrated Wolbachia pipientis in Culex pipiens and the Wolbachia species of infected members of the Aedes scutellaris group. Aposymbiosis was induced in Aedes polynesiensis by treatment with 0.017 mg/ml tetracycline hydrochloride or by rearing larvae at temperatures of 32–33°C for 5–7 days. Aposymbiotic females were incompatible with infected males but the reciprocal cross was fertile. By light and electron microscopy wolbachiae were found in ovaries of Aedes albopictus, A. cooki, A. Niuafo'ou sp., A. polynesiensis from American Samoa and Tahiti, A. riversi, A. Tafahi sp., and A. upolensis. Wolbachiae were not detected in A. alcasidi, A. oceanicus, A. triseriatus, Culex peus, two laboratory colonies of C. pipiens, Culiseta incidens, Toxorhynchites amboinensis, or T. brevipalpis. Wolbachiae were found also in the ovarian follicular epithelium of A. albopictus. This is the first record of mosquito wolbachiae in nongerminal gonadal tissue.     

Effects of acidic pH and phytoplankton on survival and condition of Bosmina longirostris and Daphnia pulex

The effects of pH, algal composition and algal biomass on abundance, size, reproduction and condition of Daphnia pulex and Bosmina longirostris were tested in a field experiment using water and natural phytoplankton assemblages from a circumneutral (pH 6.43) and a moderately acidic (pH 5.75) lake in south-central Ontario. Both species were affected by pH and phytoplankton composition, with decreased egg production, lipid reserves, body size or abundance in treatments containing algae and/or water from the more acidic lake compared to treatments containing water and phytoplankton from the circumneutral lake. This result was unexpected for Bosmina, which often increases in relative and/or absolute abundance in acidified lakes. The negative effect of acidic conditions on Bosmina suggests that the population increase observed in most acidified lakes is not due to a positive response to low pH or ambient phytoplankton, but to altered biotic interactions possibly involving reduced competition.     

Assemblages of dytiscid predators and culicid prey in relation to environmental factors in natural and clear-cut boreal swamp forest pools

Assemblages of diving beetles (Dytiscidae) and mosquito immatures (Culicidae) were studied during 1987–1988 in 40 small, more or less temporary, snowmelt pools in spruce swamp forest and clearings at the Arctic Circle in Sweden. Larger pools were warmer than smaller ones, and clearing pools were warmer than forest pools. Temperature differences between pools remained high until late July. Twenty-one dytiscid species, representing three guilds, occurred in the pools, and individual pools had 1–13 species. Ten species occurred in both habitats. A few stenotopic species of boreal swamp forests had in clearing pools apparently been replaced by some species with a preference for more productive, often man-made habitats. Six Aedes species were collected in the clearing pools. Five of these were found in the forest pools, of which three had no mosquito larvae. Dytiscid assemblages in both habitats and culicid assemblages in clearing pools showed strong nested patterns.Abundance and species richness of both culicids and dytiscids were higher in clearing than in forest pools with an area >2 m² after that the effects of pool area had been accounted for (MANCOVA). In both habitats, abundance and species richness of both culicids and dytiscids were strongly and positively correlated with a linear combination of pool area, depth and temperature (Canonical Correlation). Increasing drought frequency of pools had a negative, less significant effect on the biota. In the clearing, the abiotic correlations with abundance were somewhat weakened chiefly by the relatively low abundance values from the largest pool.Distribution and mean abundance of individual dytiscid species were positively related in the clearing pools. Flying dytiscids were trapped in the larger (1.6 m²), but not in the smaller (0.07 m²) artificial pools, and the immigration rate was markedly higher on clearings than in forest. Dug pools were colonized faster on clearings than in forest. Even the flightless Hydroporus melanarius colonized dug pools during the first year.     

Impact of chemical composition of legume residues and initial soil pH on pH change of a soil after residue incorporation

Reports on the effect of organic matter addition on soil pH have been contradictory. This study examined the effect of applying legume residues differing in concentrations of N (4.3-45.5 mg g^-1) and excess cations/organic anions (0.22–1.56 mmol g^-1) on pH change of five soils differing in initial pH (3.60–5.58 in 0.01 M CaCl₂) under sterile and non-sterile conditions. Addition of the legume residues at a level of 1% soil weight increased the pH of all soils by up to 2 units after incubation for 35 and 100 d under non-sterile conditions. Exceptions were the Lancelin (initial pH 5.06) and Kellerberin (pH 5.58) soils with addition of clover roots (excess cations 22 cmol/kg) for 100 d where soil pH decreased by 0.13–0.15 units as compared to the control. The amounts of alkalinity produced in soil correlated positively with concentrations of excess cations and total nitrogen of the added legume residues, and negatively with the initial pH of the soil. When soil was fumigated with chloroform during incubation, similar trends of soil pH changes and alkalinity production, due to legume residues addition, were displayed but the effects of the residue on alkalinity production in the Wodjil and Lancelin soils were much less than under non-sterile conditions. Direct shaking of soil with the residues under sterile conditions increased the amount of alkalinity in the soils with initial pH of 3.60–4.54, but not in the soils with initial pH of 5.06 and 5.58. The maximal alkalinity production was less than one third of that produced in the soil after 100 d of incubation under non-sterile conditions. The results suggest that the direction and the magnitude of pH change depend largely on the concentration of organic anions in the residues, initial soil pH and the degree of residue decomposition. The incorporation of crop residues, especially those with high concentrations of excess cations, is recommended in minimizing soil acidification in farming systems. This revised version was published online in June 2006 with corrections to the Cover Date.     

Interactions of intracellular pH and intracellular calcium in primary cultures of rabbit corneal epithelial cells

Summary Homeostasis of intracellular calcium ([Ca⁺⁺]_i) and pH (pH_i) is important in the cell's ability to respond to growth factors, to initiate differentiation and proliferation, and to maintain normal metabolic pathways. Because of the importance of these ions to cellular functions, we investigated the effects of changes of [Ca⁺⁺]_i and pH_i on each other in primary cultures of rabbit corneal epithelial cells. Digitized fluorescence imaging was used to measure [Ca⁺⁺]_i with fura-2 and pH_i with 2′,7′-bis(2-carboxyethyl)-5(6)-carboxyfluorescein (BCECF). Resting pH_i in these cells was 7.37±0.05 (n=20 cells) and resting [Ca⁺⁺]_i was 129±10 nM (n=35 cells) using a nominally bicarbonate-free Krebs Ringer HEPES buffer (KRHB), pH 7.4. On exposure to 20 mM NH₄Cl, which rapidly alkalinized cells by 0.45 pH units, an increase in [Ca⁺⁺]_i to 215±14 nM occurred. Pretreatment of the cells with 100 μM verapamil or exposure to 1 mM ethylene bis-(oxyethylenenitrilo)-tetraacetic acid (EGTA) without extracellular calcium before addition of 20 mM NH₄Cl did not abolish the calcium increase, suggesting that the source of the calcium transient was from intracellular calcium stores. On removal of NH₄Cl or addition of 20 mM sodium lactate, there were minimal changes in calcium even though pH_i decreased. Treatment of CE cells with the calcium ionophores, ionomycin and 4-bromo A23187, increased [Ca⁺⁺]_i, but produced a biphasic change in pH_i. Initially, there was an acidification of the cytosol, and then an alkalinization of 0.10 to 0.11 pH units above initial values. When [Ca⁺⁺]_i was decreased by treating the cells with 5 mM EGTA and 20 μM ionomycin, pH_i decreased by 0.35±0.02 units. We conclude that an increase in pH_i leads to an increase in [Ca⁺⁺]_i in rabbit corneal epithelial cells; however, a decrease in pH_i leads to minor changes in [Ca⁺⁺]_i. The ability of CE cells to maintain proper calcium homeostasis when pH_i is decreased may represent an adaptive mechanism to maintain physiological calcium levels during periods of acidification, which occur during prolonged eye closure.     

Effects of positive interactions, size symmetry of competition and abiotic stress on self-thinning in simulated plant populations

Background and Aims

Competition drives self-thinning (density-dependent mortality) in crowded plant populations. Facilitative interactions have been shown to affect many processes in plant populations and communities, but their effects on self-thinning trajectories have not been investigated.

Methods

Using an individual-based ‘zone-of-influence’ model, we studied the potential effects of the size symmetry of competition, abiotic stress and facilitation on self-thinning trajectories in plant monocultures. In the model, abiotic stress reduced the growth of all individuals and facilitation ameliorated the effects of stress on interacting individuals.

Key Results

Abiotic stress made the log biomass – log density relationship during self-thinning steeper, but this effect was reduced by positive interactions among individuals. Size-asymmetric competition also influenced the self-thinning slope.

Conclusions

Although competition drives self-thinning, its course can be affected by abiotic stress, facilitation and competitive symmetry.     

Effects of aluminum,calcium and low pH on egg hatching and nymphal survival of Cloeon triangulifer McDunnough (Ephemeroptera: Baetidae)

Laboratory experiments were conducted to assess the effects of aluminum, calcium and low pH on egg hatching and nymphal survival of the mayfly Cloeon triangulifer. Percent successful hatch (living nymphs breaking free of the chorion) decreased and percent partial hatch (nymphs dying attached to the chorion) increased with increasing acidity (pH 7.5–3.0). Most hatches occurring below pH 5.0 were partial hatches. Decreased time of exposure to acidic waters increased percent successful hatch and decreased percent partial hatch. Time to first hatch was not affected by pH. Eggs were incubated in acidic waters (pH 4.0 and 5.5) with additions of calcium (10 and 100 mg l^–1) and aluminum (100 and 500 g l^–1). Aluminum decreased percent successful hatch and increased percent partial hatch and calcium increased both percent successful hatch and percent partial hatch. Time to first hatch was increased by both aluminum and calcium. The 96 h LC50 for small nymphs was pH 4.75. Addition of aluminum (100 and 500 µg l^–1) to acidic waters (pH 4.0 and 5.0) reduced nymphal mortality by 8–22%. Toxic effects of low pH on egg hatching and early nymphs may contribute to the absence of mayflies from acidified habitats.Contribution No. 1469 of the Maine Agricultural Experiment Station, University of Maine, Orono, Maine 04469 USA.Contribution No. 1469 of the Maine Agricultural Experiment Station, University of Maine, Orono, Maine 04469 USA.     

Structure of host-odour plumes influences catch of Anopheles gambiae s.s. and Aedes aegypti in a dual-choice olfactometer

The effects of structure, concentration and composition of host‐odour plumes on catch of female Anopheles gambiae Giles sensu stricto and Aedes aegypti (L.) (Diptera: Culicidae) were investigated in a dual‐choice olfactometer. We demonstrate that the fine‐scale structure of host‐odour plumes modulates capture of An. gambiae and Ae. aegypti. In both species homogeneous skin‐odour plumes result in trap entry, whereas homogeneous CO₂ plumes reduce trap catch. Reduced trap catch also result from combining skin odour with a homogeneous CO₂ plume. Trap capture rates in homogeneous CO₂ plumes were concentration‐dependent and differed between the two species. Electric nets placed in front of the trap entrances intercepted mosquitoes before they could enter the traps. This showed that An. gambiae flew along CO₂ plumes, but did not enter the traps. Survivorship analysis of the trap‐entry times of Ae. aegypti indicated interactions between the time until capture and treatment. The assay's duration therefore can alter the distribution in a dual‐choice olfactometer.     

Consumer body composition and community structure in a stream is altered by pH

1. Low pH inhibits microbial conditioning of leaf‐litter, which forms the principal energy input to many headwater streams. This reduces food quality and availability for the shredder assemblage, thereby creating a potential bottleneck in the flux of energy and biomass through acidified food webs. 2. We explored the consequences of acidity on the well‐characterised community of Broadstone Stream in southeast England, by quantifying the physiological condition (protein and lipid content) of three dominant shredder species (Leuctra nigra, L. hippopus and Nemurella pictetii) and relating this to changes in the numerical abundance and biomass of invertebrates across a longitudinal pH gradient (5.3–6.5). 3. Total taxon richness increased with pH, as did shredder diversity. The acid‐tolerant stonefly, L. nigra, exhibited a positive correlation between pH and protein content, but its abundance was suppressed in the less acid reaches. These results suggest that the impacts of environmental stressors might be manifested differently at the population (i.e. numerical and biomass abundance) versus the physiological (i.e. protein content of individuals) levels of organisation. Body composition of L. hippopus and N. pictetii did not exhibit any significant relationship with stream pH in the field. 4. The survey data were corroborated with a laboratory rearing experiment using N. pictetii, in which survival rate, growth rate, and protein and lipid content of individuals were measured in stream water of differing pH and acid versus circumneutral microbial conditioning regimes. Acid‐conditioned leaves were associated with increased mortality and reduced protein content in consumers’ tissues, with acid water also having the latter effect. 5. Our results suggest that biochemical constraints within key taxa might create energy flux bottlenecks in detrital‐based food webs, and that this could ultimately determine the productivity of the entire system. Hence assays of the body composition of macroinvertebrates could be an effective new tool that complements population level studies of the impacts of stressors in fresh waters.     

Effects of experimental acidification on a lotic macroinvertebrate community

A three month experimental acidification was carried out on lotic bottom communities. Experiments were conducted under semi-natural conditions in plasticized wooden channels. Acidified communities (pH 4.0), with or without added aluminum, were compared with a reference community (pH 6.3–6.9). Added aluminum concentrations were respectively 0.2 and 0.4 mg 1^–1 in experiments performed in 1982 and 1983. Water chemistry and taxonomic composition of the macroinvertebrate communities were monitored. Under acidified conditions, results were similar, with or without added aluminum. Mean abundances of all groups of organisms were lowered. Mayflies nearly completely disappeared from the acidified channels. The only organism not affected by the acidification was Microtendipes sp. Differences in the organism response were observed: Orthocladiinae (Rheocricotopus, Parametriocnemus, Corynoneura, Thienemanniella, Nanocladius, Cricotopus) and Ephemeroptera (Baetis, Habrophlebia, Habrophlebiodes, Paraleptophlebia, Ephemerella), especially early instars, were very sensitive to low pH, Chironomini and Tanypodinae were much less sensitive, while Tanytarsini were intermediate; Oligochaeta and Nematoda were difficult to classify, their response being different from one year to another. Organisms inhabiting the surface of artificial substrates disappeared very rapidly from the system, while those buried inside had a delayed reaction to acidification. Aluminum which was mainly in the monomeric form was not responsible for community modifications. Direct action of hydrogen ions through a physiological stress seems a more credible explanation. These results, induced by a continuous experimental acidification, suggest that if this small headwater stream undergoes acidification, the resulting invertebrate community will be very simplified, with only resistant species able to cope with the acid conditions.     

Hemolymph pH of the larvae of three species of mosquitoes,and the effect of Romanomermis culicivorax parasitism on the blood pH of Culex pipiens

The hemolymph pHs of late fourth-instar Culex pipiens, Aedes taeniorhynchus, and Toxorhynchites amboinensis were measured with pH-sensitive glass microelectrodes and were slightly alkaline; pH 7.51, 7.62, and 7.37, respectively. The hemolymph pH remained relatively constant during the development of C. pipiens larvae through the third and early and late fourth instars. The hemolymph pH in C. pipiens larvae parasitized with the mermithid nematode Romanomermis culicivorax was unaltered. These measurements provide an approximate pH (ca 7.4) which is normal for the hemolymph of larval mosquitoes, and should be useful for further development of a culture medium for R. culicivorax.