Predator-prey interactions and community structure: chironomids,mosquitoes and copepods in Heliconia imbricata (Musaceae)

Summary Evidence from both field observations and experimental work indicates that predation by larvae of a midge, Pentaneura n. sp. (Chironomidae), causes the low densities of mosquito larvae (Culicidae) found in the water filled bracts of Heliconia imbricata (Musaceae), microhabitats typically colonized by mosquitoes. This predation affects 2 species of mosquitoes, Wyeomyia pseudopecten, a resident species, and Trichoprosopon digitatum, a non-resident species. Predation keeps resident mosquito densities low while completely excluding the nonresident mosquito from the habitat. Both these effects of predation depend on the presence of an abundant alternative prey, an undescribed species of harpacticoid copepod found in the bracts. These copepod prey sustain chironomids when resident mosquito densities are low, permiting predator densities to remain high enough to exclude the non-resident mosquito. I discuss the evolutionary and ecological implications of predation structuring communities.     

Understanding gregariousness in a larval Lepidopteran: the roles of host plant, predation, and microclimate

Abstract.  1. Many moth and butterfly larvae are gregarious early in development, but become solitary in late instars. This ontogenetic variation in behaviour is probably the result of temporal changes in the costs and benefits associated with gregariousness. This study provides observational and experimental evidence that, in one particular moth species, a series of different ecological factors influence larval behaviour at different times during development.
2. Field observations show that young caterpillars of the limocodid Doratifera casta form large aggregations while foraging, but that mature larvae are largely solitary.
3. A field experiment revealed that individual first to third instar larvae in larger groups develop more rapidly, but that group size had no detectable influence on survival. The developmental advantage associated with gregariousness is affected by host plant species, but not by predator exclusion, suggesting that group living in these cryptic early instar larvae promotes feeding facilitation, but does not provide individuals with protection from natural enemies.
4. Laboratory experiments revealed that aposematic fourth instar caterpillars in large groups were less likely to be attacked by a generalist insect predator than those in small groups.
5. Field observations provided no evidence that group living affects body temperature, suggesting that microclimatic factors do not favour gregariousness in this species.
6. It is concluded that gregariousness in D. casta confers at least two different advantages on larvae at different stages early in development, but that these advantages disappear, or are outweighed by costs associated with intraspecific competition, in final instars.     

Selective and frequency dependent predation of aquatic mosquito predator Diplonychus indicus Venkatesan & Rao (Hemiptera: Belostomatidae) on immature stages of three mosquito species

Frequency dependent mosquito larval size (II and IV instars) and species selection by the water bug Diplonychus indicus against three mosquito species Culex quinquefasciatus, Aedes aegypti and Anopheles stephensi was studied in the laboratory. The different frequencies used for each species selection were 20:30:50, 30:50:20, 50:20:30, 25:35:40, 35:40:25 and 40:25:35 of fourth instars of the respective three prey species. All nymphal water bugs (I–V instars) selected IV instar mosquito larvae and the mean proportion of late (larger) larvae eaten by the predator instars was significantly higher than the mean proportion of early (smaller) larvae eaten (F= 2.28; P < 0.001). In all six ratios used to determine the frequency dependent mosquito species selection, all the stages of the water bug selected Ae. aegypti over the other two species (F= 452.43; P < 0.001). The mean number of mosquito larvae eaten increased as its density increased based on various ratios of larvae offered. The study indicated that the predatory efficiency of D. indicus was high when Ae. aegypti was offered as prey, suggesting the utility of this mosquito predator in the control of dengue vectors.     

Regulation of host diapause by an insect parasitoid

Abstract. 1. The interaction between larval development and parasitism by the braconid wasp Cotesia koebelei (Riley), was investigated in a population of the butterfly Euphydryas editha (Boisduval) (Nymphalidae). In this population, the butterfly host has an obligatory overwintering larval diapause.
2. It was found that E. editha larvae harbouring parasitoids were more likely to pass through an extra feeding instar before entering diapause than were non-parasitized conspecifics.
3. In addition, some individuals that were experimentally exposed to multiple parasitoid attacks bypassed diapause completely; these larvae passed through five or six feeding instars, reaching sizes typical of final instar post-diapause larvae.
4. The observed effect of superparasitism occurred regardless of whether the host larvae subsequently produced mature parasitoids, suggesting that parasitoid attack is sufficient to invoke the response.
5. It is proposed that the parasitoid C.koebelei regulates the number of pre-diapause feeding instars of its insect host E. editha, and that some component of the female venom, injected at oviposition, is responsible for this regulation.     

Food selection of Tanytarsus gracilentus larvae (Diptera: Chironomidae): An analysis of instars and cohorts

Gut contents of the detritivorous Tanytarsus gracilentus larvae were studied at one site in Lake Myvatn, Iceland. Samples were collected with a sediment corer. Sediment samples were taken from the surface of the intact core before the larvae were collected. Gut contents were compared between different larval instars and cohorts, and to the surface sediment. The composition of the gut contents of all instars was different from that of the sediment surface. The first instar selected small diatoms of the genus Fragilaria, as single and paired cells, and the fourth instar selected detritus items of more variable size and origin. Apart from the third instar larvae, there were similar trends in food selectivity between the same instar of different cohorts. No major changes occurred in the composition of the surface sediment during the time of this study.     

Parasitoid-induced behavioral alterations of Aedes aegypti mosquito larvae infected with mermithid nematodes (Nematoda: Mermithidae).

A wide range of parasites are known to cause behavioral changes in their hosts and parasitized insects are especially amenable to the study of such changes. The majority of studies addressing parasite-induced behavioral alterations have focused on parasites with complex life cycles and the adaptive nature of such changes. Behavioral changes caused by parasitoids, single-host parasites that kill their host upon emergence, have been studied less and the adaptive nature of these changes is likely to be different than those in complex life cycles. I investigated behavioral alterations in Aedes aegypti mosquito larvae infected with parasitoid nematodes (family Mermithidae). I conducted several experiments in which I tested the following hypotheses: 1) Mermithid nematodes induce behavioral changes in mosquito larvae and the changes are density dependent. 2) Different species of mermithid nematodes induce similar changes in mosquito larvae behavior. 3) Behavioral alterations vary with mermithid developmental stage. 4) Mosquito larvae infected with mermithid nematodes behave similarly to uninfected food-deprived mosquito larvae. I found that 4th instar Ae. aegypti infected with Romanomermis culicivorax or Strelkovimermis spiculatus exhibited resting behaviors significantly more often than uninfected controls but that intensity of infection did not affect activity levels. In earlier instars, infected mosquito larvae were more active than uninfected control larvae in some behaviors associated with feeding. There was no significant difference between infected and uninfected food-deprived mosquitoes in nine of the ten behaviors observed. The decrease in activity of late instar Ae. aegypti larvae infected with mermithids may be a parasitoid adaptation that reduces the risk of predation and thus increases host and parasitoid survival. The increase in feeding activity in earlier instars as well as the similarity between uninfected food-deprived and infected Ae. aegypti behavior may indicate that these behaviors are adaptive for the parasitoid, increasing nutritional acquisition for successful parasitoid development.     

Mechanical limitations on feeding by meadow spittlebugs Philaenus spumarius (Homoptera: Cercopidae) on wild and cultivated host plants

Abstract. 1. Field observations on the distribution of Philaenus spumarius (L.) on pearly everlasting (Anaphalis margaritacea) and alfalfa (Medicago sativa) showed that the range of feeding sites exploited increased with instar. Development of trichomes, depth of xylem elements, and tissue hardness were measured as possible factors determining the suitability of sites on food plants.
2. Spittlebugs caged outside their normal feeding sites experienced difficulty feeding. Experimental removal of trichomes showed that trichomes prevented younger instars from feeding on the stem. The depth of the xylem elements and tissue hardness were not significant barriers to feeding near the stem apex.
3. By caging fourth and fifth instar nymphs at increasing distances below the terminal bud on A.margaritacea stems, we showed feeding ability declined with distance below the apex, and feeding was more restricted in fourth compared to fifth instars. A similar decline was demonstrated in experiments with fifth instars on M.sativa.
4. On A.margaritacea, this restriction was associated with increasing tissue hardness of the stem with distance from the apex and the presence of trichomes in regions with hard tissues. On M.sativa the associated barriers were a combination of tissue hardness and reduction in the availability of xylem vessels in the penetrable region of the stem. Although the tissues impeding stylet penetration differed in the two hosts, the critical factor in both species was the increase in the amount of lignification of those tissues at increasing distances from the terminal bud.     

Interactions with native mosquito larvae regulate the production of Aedes albopictus from bromeliads in Florida

Abstract. 1. Immatures of the invasive container mosquito Aedes albopictus occur in water-holding tanks and axils of ornamental bromeliads in Florida, where this species established and became abundant in the late 1980s and early 1990s.
2. The numbers of A. albopictus in bromeliads in gardens vary geographically in Florida, being significantly higher in northern sites where two species of Wyeomyia mosquitoes, native specialists of this microhabitat, are absent.
3. Causes of the negative relationship between abundances of A. albopictus and Wyeomyia spp. were investigated experimentally using Billbergia pyramidalis , an introduced bromeliad popular in Florida gardens.
4. Egg laying by A. albopictus in B. pyramidalis confined in a large outdoor cage was unaffected by the presence or absence of Wyeomyia spp. fourth instars, indicating that immatures of the native bromeliad mosquito species do not deter oviposition by the invasive species in this microhabitat.
5. Growth and survivorship of A. albopictus first instars in tanks of B. pyramidalis were negatively affected by the presence of fourth instar, but not first instar, Wyeomyia spp. larvae at natural densities. Stimulative effects of leaf litter on A. albopictus growth and survivorship in this microhabitat were independent of the presence or absence of Wyeomyia spp. immatures.
6. These results demonstrate that interspecific larval interactions regulate the production and explain the geographic patterns in abundance of A. albopictus from ornamental bromeliads in Florida.     

Distribution patterns of hummingbird flower mites (Gamasida: Ascidae) in relation to floral availability on Heliconia inflorescences

I analyzed the dispersion patterns of ascid mites (Gamasida:Ascidae) on inflorescences of Heliconia trinidatis in relationto the temporal and spatial distribution of open flowers. Openflowers last only a few hours and are the locations of foodas well as the sole sites from which mites disperse on visitinghummingbirds. All inflorescences were inhabited by populationsof nectarivorous Rhinoseius trinitatis and omnivorous Lasioseiuselegants. All demographic groups of R. trinitatis were associatedstrongly with bracts bearing open flowers on the day of inflorescencecollection or with bracts that would have had an open floweron the day after collection. For L. elegans, only larval distributioncoincided with open flowers. Laboratory observations demonstratedthat mites move throughout an inflorescence at night and preferentiallyaggregate where the next flower will open. Mites enter flowersimmediately at, or just before, anthesis, and unerringly exitopen flowers well in advance of floral abscission. Hummingbirdflower mites exhibit a variety of behavioral adaptations, indicatingextraordinary sensitivity to the physiological and biochemicalnuances of their host inflorescences.     

Stage-dependent predation on competitors: consequences for the outcome of a mosquito invasion

1. Predator-mediated coexistence occurs when predation allows competitors to coexist, due to preferential consumption of a superior competitor relative to an inferior competitor. Differences between the native treehole mosquito ( Aedes triseriatus ) and the co-occurring Asian tiger mosquito ( Aedes albopictus ) in anti-predatory larval behaviours account, in part, for the greater vulnerability of this invasive species to native predatory midge ( Corethrella appendiculata ). We test the hypothesis that stage-dependent differences in the sizes of A. albopictus and A. triseriatus larvae, relative to the size-limited C. appendiculata , contribute to differential consumption and the likelihood of predator-mediated coexistence of these competitors.
2. In all instars, larvae of A. triseriatus were larger than A. albopictus of the same stage. Third and fourth instar C. appendiculata selectively consumed late-stage A. albopictus in preference to same-stage A. triseriatus . Small, early-stage prey larvae did not differ in vulnerability to predation, but large, late-stage larvae differed significantly in vulnerability to predation, probably owing to size-limited predation by fourth instar C. appendiculata. This effect was less pronounced for third instar C. appendiculata .
3. Prey size, in conjunction with anti-predatory behavioural responses, alters the probability of predator-mediated coexistence. A stage-structured predation model showed that equally vulnerable early stages reduce the range of environmental conditions (productivities) in which predator-mediated coexistence is possible, increasing the likelihood of both competitive exclusion of the resident species or failure of the invasive to establish. These results underscore the importance of stage-dependent interspecific differences in predator–prey interactions for determining how predators may affect community composition.     

Unsuitability of Cnesterodon decemmaculatus (Jenyns, 1842) for mosquito control in Uruguay: evidence from food‐preference experiments

We analyzed the feeding preference of Cnesterodon decemmaculatus, a small‐bodied poecilid native from the Rio de la Plata and proximate Atlantic Basins in South America. This species has a wide distribution in Uruguayan water bodies but its effectiveness as a predator of mosquito larvae has not been tested. In laboratory trials, five aquatic invertebrates were offered simultaneously as potential prey to fish: Daphnia pulex (Cladocera), copepods, two different instars of mosquito larvae (Culex pipiens), and the 4^th instar of Chironomidae larvae. Preference was measured by the Chesson's electivity index (α). In order to determine differences in prey preference according to fish size, individuals ranging from 9.5 mm to 35.3 mm were classified in three different body size classes: small, medium, and large. Small fish showed preference for copepods, while medium‐sized fish preferred the smallest mosquito larvae instars and Chironomidae larvae. We conclude that C. decemmaculatus is a zooplankton facultative‐feeder fish that prefers large‐bodied zooplankton but is a weak predator of mosquito larvae. Thus, the introduction of C. decemmaculatus as a biological‐control agent in natural environments is not an effective strategy.     

Asymmetric intraspecific competition among larvae of the damselfly Ischnura elegans (Zygoptera: Coenagrionidae)

Abstract. 1. A laboratory competition experiment is described in which the growth and development rates of larvae of the damselfly Ischnura elegans (Lind.) were measured over an entire instar.
2. Two larval instars which commonly occur together in the field were used in the experiment; they were maintained with a superabundance of prey and either larvae from the same or the larger/smaller instar.
3. Small larvae suffered increased development times and decreased size increases at the moult in the presence of large larvae but similar interference effects were not evident when these smaller larvae were in the presence of other small larvae.
4. Development time and size increases of large larvae were not significantly affected by the presence of small larvae.
5. Irrespective of the instar combinations investigated, interference effects were reduced when there were more perches available, although in only a few cases was this reduction significant.
6. The consequences of the asymmetric competition reported in the experiment for the study of lifetime reproductive success in damselflies are discussed. Late emerging adults may incur reduced reproductive success.     

Achieving high sexual size dimorphism in insects: females add instars

Abstract.  1. In arthropods, the evolution of sexual size dimorphism (SSD) may be constrained by a physiological limit on growth within each particular larval instar. A high SSD could, however, be attained if the larvae of the larger sex pass through a higher number of larval instars.
2. Based on a survey of published case studies, the present review shows that sex-related difference in the number of instars is a widespread phenomenon among insects. In the great majority of species with a sexually dimorphic instar number, females develop through a higher number of instars than males.
3. Female-biased sexual dimorphism in final sizes in species with sexually dimorphic instar number was found to considerably exceed a previously estimated median value of SSD for insects in general. This suggests a causal connection between high female-biased SSD, and additional instars in females. Adding an extra instar to larval development allows an insect to increase its adult size at the expense of prolonged larval development.
4. As in the case of additional instars, SSD is fully formed late in ontogeny, larval growth schedules and imaginal sizes can be optimised independently. No conflict between selective pressures operating in juvenile and adult stages is therefore expected.
5. In most species considered, the number of instars also varied within the sexes. Phenotypic plasticity in instar number may thus be a precondition for a sexual difference in instar number to evolve.     

Predation by Halmus chalybeus (Coleoptera: Coccinellidae) on Ceroplastes destructor and C. sinensis (Hemiptera: Coccidae: Ceroplastinae) Infesting Citrus in Northland, New Zealand

Predation by Halmus chalybeus (steelblue ladybird) on two species of wax scale was studied on citrus orchards in Northland, New Zealand. Field experiments using an exclusion technique of enclosing citrus branches in bags, found that larval and adult H. chalybeus preyed on first and second instars of both Ceroplastes destructor (white wax scale) and C. sinensis (Chinese wax scale), but not third instar C. destructor. Third instar C. sinensis and adults of both species were not tested but are rarely, if ever, attacked by H. chalybeus. The sampling of scale and ladybird populations and field experiments showed that few scales of either species survived past the second instar stage where H. chalybeus was numerous. The feeding rate of H. chalybeus on settled first and second instar scales was estimated inside bagged branches. Adults consumed on average 15.6 C. destructor and 13.3 C. sinensis per day per ladybird, while larvae ate 9.7 C. destructor per day. These feeding rates accounted for the experimental reduction of scale populations. H. chalybeus is a useful natural enemy for the control of C. destructor and C. sinensis when its activity is not disrupted by pesticides.     

Intraspecific competition amongst larval Aedes aegypti: food exploitation or chemical interference?

Abstract. 1. Competition within and between the larval instars of the yellow fever mosquito, Aedes aegypti , can be measured by its effect on stage duration. In a series of laboratory experiments the relative importance of chemical interference and food exploitation in mediating competition between first and fourth instar larvae was investigated.
2. In contrast to the results of three previous studies, I found no evidence that a chemical growth retardant played any part in greatly prolonging the stage durations of larvae in both age classes.
3. When competition between the two age classes became important, the relative increase in stage duration wasgreater for first instar larvae than for fourth instar larvae. This result can be reproduced with Gilpin & McClelland's (1979) model of competition by food exploitation alone, providing the range of available food particle size is assumed to be an increasing function of age. An additional, but less important, refinement makes the conversion efficiency of food into larval biomass a decreasing function of age.     

Snail predation by larvae of Sepedon scapularis Adams (Diptera: Sciomyzidae), a potential biocontrol agent of snail intermediate hosts of schistosomiasis in South Africa

Malacophagous larvae of the fly Sepedon scapularis Adams were shown experimentally to be effective predators of three species of aquatic pulmonate snails tested as prey: Bulinus africanus (Krauss) an important intermediate host of Schistosoma haematobium (Bilharz), Bulinus tropicus (Krauss) and the invasive species Physa acuta Draparnaud. Survival of S. scapularis larvae from instar to instar was negatively affected by the size of prey snails, since larvae tended to be asphyxiated by the mucous secretions of the snails, or by the larval hydrofuge hairs becoming entangled in snail faeces. In experiments to test the choice of S.scapularis larvae for different species and sizes of snails, B.africanus was significantly disfavoured compared with the other two snail species. Small snails (< 3 mm) were more frequently killed by all three larval instars of S.scapularis. First instars killed few, if any, large snails (> 7 mm), whereas second and third instars preyed effectively on all sizes of snails. Third instars killed significantly more snails than younger instars. For larvae offered only one species of snail as prey, the mean total number of snails killed per larva during its entire development was 49 B.africanus, 45 B.tropicus or 34 P.acuta. It is concluded that S.scapularis is potentially useful as a biological control agent for use against indigenous Bulinus and exotic Physa snails.     

Reproductive biology of Syagrus coronata (Arecaceae): sex‐biased insect visitation and the unusual case of scent emission by peduncular bracts

• Several monoecious species of palms have developed complex strategies to promote cross‐pollination, including the production of large quantities of floral resources and the emission of scents that are attractive to pollinators. Syagrus coronata constitutes an interesting model with which to understand the evolution of plant reproductive strategies in a monoecious species adapted to seasonally dry forests.
• We monitored blooming phenology over 1 year, during which we also collected and identified floral visitors and putative pollinators. We identified potential floral visitor attractants by characterizing the scent composition of inflorescences as well as of peduncular bracts, during both male and female phases, and the potential for floral thermogenesis.
• Syagrus coronata produces floral resources throughout the year. Its inflorescences are predominantly visited by a diverse assortment of small‐sized beetles, whose richness and abundance vary throughout the different phases of anthesis. We did not find evidence of floral thermogenesis. A total of 23 volatile compounds were identified in the scent emitted by the inflorescences, which did not differ between male and female phases; whereas the scent of the peduncular bracts was composed of only 4‐methyl guaiacol, which was absent in inflorescences.
• The composition of floral scent chemistry indicates that this palm has evolved strategies to be predominantly pollinated by small‐sized weevils. Our study provides rare evidence of a non‐floral scent emitting structure involved in pollinator attraction, only the second such case specifically in palms. The peculiarities of the reproductive strategy of S. coronata might play an important role in the maintenance of pollination services and pollen dispersion.

     

Dropping behaviour of larvae of aphidophagous ladybirds and its effects on incidence of intraguild predation: interactions between the intraguild prey, Adalia bipunctata (L.) and Coccinella septempunctata (L.), and the intraguild predator, Harmonia axyridis Pallas

Abstract.  1. Two experiments were performed in the laboratory to assess the behaviour of dropping from a host plant as a defence against intraguild predation in aphidophagous ladybird larvae.
2. In the first experiment, encounters were observed on bean plants between fourth instars of the intraguild predator species, Harmonia axyridis , and first instars of two other ladybird species, Adalia bipunctata (L.) and Coccinella septempunctata (L.). The percentages of first instars of the latter two species that dropped from the plant in response to attack differed dramatically, with 47.5% of C. septempunctata first instars dropping vs. 0% of A. bipunctata .
3. In the second experiment, first instars of A. bipunctata or C. septempunctata and a fourth instar of H. axyridis were allowed to forage together on bean plants for 3 h. During this time, 44.3% of C. septempunctata larvae dropped from the plant, but less than 2% of A. bipunctata larvae did so. In contrast, 95.0% of A. bipunctata larvae fell victim to intraguild predation by H. axyridis vs. only 54.5% of C. septempunctata larvae.
4. The significance of dropping behaviour of ladybird larvae as a defence against intraguild predation, and the relationship of dropping behaviour to species-specific habitat affinity of ladybirds, is discussed.     

Studies on the biology of Chaoborus flavicans (Meigen) (Diptera: Chaoboridae) in a fish-free eutrophic pond,Japan

Population dynamics of Chaoborus flavicans larvae of various instars was studied from November 1986 to December 1987 in a eutrophic, fish-free pond, Japan. First and 2nd instar larvae were observed from late April to late October, indicating a reproductive period of about half a year. C. flavicans overwintered in the 4th instar larvae. In water column samples, total density of all instars was 680–23680 m^-2, and pupal density 0–2600 m^-2; larvae of the 1st, 2nd, and 3rd instars showed 5–6 density peaks in 1987, suggesting that 5–6 generations occur during a year (peaks of the 4th instar larvae were not clear, probably due to their longer development than those of younger instars). In sediment samples, no 1st and 2nd instar larvae were found, 3rd instar larvae were found occasionally but density of the 4th instar larvae was 280–18600 m^-2, and pupal density varied between 0–502 m^-2. Fouth instar larvae accumulated in sediment in the cold season and in the water column in the warm season; high temperature and low oxygen concentration were the most important factors limiting the distribution of larvae in the sediment in summer in the NIES pond. The dry weight of total C. flavicans larvae was 0.08–4.2 g m^-2 in sediment samples and 24–599 μg l^-1 (0.10–2.40 g m^-2) in water column samples. Comparisons of maximum densities in the NIES pond in different years and in waters of different trophic status show that density is generally higher in eutrophic than in oligotrophic habitats. This revised version was published online in July 2006 with corrections to the Cover Date.     

Experimental study of larval efficiency of Gambusia affinis holbrooki (GIRARD, 1859) (fish-Poecilidae)

Experimental study of efficiency of Gambusia affinis holbrooki (GIRARD, 1859) (Fish-Poecilidae). Gambusia affinis is a culiciphage fish which was introduced in Tunisia in 1929 to control mosquito larvae. In the present study, we confirm, under laboratory conditions, the good efficacy of Gambusia affinis against mosquito larvae. The consumption of larvae decreases with the evolution of larvae instars and increases, for the same instar, in relation with the fish size. In fact, the size of the prey is related to the predator's size: fish belonging to the length class [20-24 mm] prefer IIIrd instar, fish from class length [25-29 mm] prefer pupae and IVth instar. The trophic test shows that G. affinis has a clear preference for Culicidae larvae, except fish belonging to the 2 length classes [35-39 mm] and [40-44 mm] for which Culicidae and Ephemeroptera have similar preference. In addition, at high temperature (30 degrees C), Gambusia affinis has a higher efficiency.