The involvement of sand disturbance, cannibalism and intra-guild predation in competitive interactions among pit-building antlion larvae

Competition in trap-building predators such as antlion larvae is a complex biotic interaction, potentially involving exploitation competition, sand throwing (i.e., interference competition), cannibalism and intra-guild predation. We investigated the short-term behavioral and developmental responses of the strict sit-and-wait antlion predator Myrmeleon hyalinus to sand disturbance (i.e., quantification of the impact of severe sand throwing), and to con- and hetero-specific competition by a larger sit-and-pursue antlion species Lopezus fedtschenkoi. We found that antlions subjected to sand disturbances reduced their pit construction activity and relocated less often. Furthermore, the reduction in pit construction activity was stronger among antlions subjected to disturbances prior to feeding. Almost no death occurred during the sand disturbance experiment, but as expected, disturbances caused reductions in the relative growth rates of antlions. This negative effect was stronger in the group exposed to sand disturbances prior to feeding. The presence of the sit-and-pursue competitor led to reductions both in pit construction and in relocation activities of M. hyalinus. Although the per-capita food supply was identical in both experiments, only 48% of M. hyalinus larvae survived the competition experiment, and this pattern was consistent between the con- and hetero-specific treatments. However, in the presence of hetero-specific competitors, the relative growth rate of surviving larvae was significantly lower than that measured in the presence of con-specific competitors. Our study demonstrates that investigating the different components of complex biotic interactions can markedly improve our understanding of how these different factors interact to influence the behavior and life history of organisms.     

Foraging in the ant-lionMyrmeleon mobilis hagen 1888 (neuroptera: Myrmeleontidae): Behavioral flexibility of a sit-and-wait predator

The relative effects of foraging benefits (food intake) and costs (disturbance = pit destruction) on growth rate and foraging behavior of larvae of the ant-lionMyrmeleon mobilis were investigated. In a laboratory experiment second-instar larvae were subjected to combinations of three rates of feeding and three rates of disturbance. Pit relocations were generally rare but occurred more often in starved larvae. Disturbance rate had no significant effect on pit relocation rate. Feeding rate was a major determinant of the energy allocated in pit construction and maintenance (positive relationship). In a second experiment third-instar larvae encountered changing rates of disturbance. Pit size was reduced as a response to increased rates of disturbance. Disturbance had no detectable effect on growth. In general,M. mobilis larvae were cost-conservative, foragers.     

The effects of starvation and repeated disturbance on mass loss,pit construction,and spatial pattern in a trap‐building predator

1. Starvation tolerance is an important trait for animals, as most will encounter starvation within their lifetime. Sit‐and‐wait predators are better adapted to starvation owing to their naturally low encounter rate with prey. 2. Starvation tolerance was studied under three levels of disturbance of wormlion larvae, a strict sit‐and‐wait predator that constructs pits. 3. Frequently disturbed wormlions constructed pits less often, and larger individuals continued to construct pits more frequently than smaller ones. It was expected that a high disturbance level would lead to a high rate of mass loss, however, surprisingly, the rate of mass loss was not higher for the frequently disturbed group. This suggests that the energetic cost of pit construction and maintenance is not as high as previously suggested for other pit‐building predators. 4. Larger individuals tolerated starvation better, in losing a lower proportion of their initial body mass and having higher chances of survival throughout the experiment. 5. The effect of starvation on the distance to neighbours was also investigated, and starved individuals were expected to maximise this distance in order to avoid interference competition. However, wormlions were usually clumped, and starvation or feeding had no effect on the pits' spatial pattern, suggesting that interference competition plays a minor role in this species. 6. Generally, wormlion larvae demonstrated a high starvation tolerance and low mortality rates even after 9 weeks of starvation.     

Dispersal between host populations in field conditions: navigation rules in the parasitoid Venturia canescens

Abstract. 1. Dispersal is a life-history trait that can have great ecological and evolutionary consequences, however understanding of how insects disperse is limited.
2. Navigation rules of the solitary koinobiont parasitoid of the pyralid moth larvae Venturia canescens (Gravenhorst) were studied in conditions that it is likely to meet when dispersing between host populations and in the absence of cues related directly to the presence of hosts.
3. Mark–release–recapture experiments were conducted in a natural host-free habitat, and letting the animals disperse for different periods.
4. In the presence of vegetation, wasps seemed to disperse rapidly (1 h for an area of ≥ 1 ha) and capture rates were independent of both dispersal time and distance from the release point.
5. The navigation rules of V. canescens during dispersal between tree stands can be summarised as: move up- or down-wind, avoid or pass through open, sunny areas, and go for shady and dense vegetation.
6. The consequences of the navigation rules for host–parasitoid dynamics are discussed in relation to different spatial scales.     

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.     

Roles of endothermy in niche differentiation for ball-rolling dung beetles (Coleoptera: Scarabaeidae) along an altitudinal gradient

Abstract.  1. An analysis of whether niche differentiation in ball-rolling dung beetles can be explained by the way in which they regulate their body temperature was conducted.
2.  A priori assumptions were: (i) if thermoregulation affects niche partitioning, sympatric species must have different endothermic strategies that minimise encounters; or, alternatively (ii) if two co-occurring species show the same thermoregulation pattern and their flight periods overlap, they might be avoiding competition by exhibiting different resource preferences or different food relocation behaviour.
3. The ball-rolling dung beetles studied showed a hierarchical structure based on the species' endothermic capacity, measured as temperature excess [ T _ex= difference between body temperature ( T _b) and ambient temperature ( T _a)]. Those with a high T _ex (10–15 °C) were located exclusively at altitudes >1000 m a.s.l. On the coastal plains, species with a high T _ex were restricted to flying at night when the T _a was lower. Species with a lower T _ex (less than 10 °C higher than T _a) were found in the coastal plains zone.
4. Where there was sympatry with similar trophic habits, the species involved showed very different thermal niches, and where there was significant overlap of thermal niches between sympatric species, trophic habits of species were very different.
5. The results suggest that it is possible to use the concept of the thermal niche as a tool to explain interspecific interactions and the spatial distribution of species.     

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.     

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.     

Intrinsic competition between larval parasitoids with different degrees of host specificity

Abstract.  1. Interspecific competition among parasitoids may play a key role in the community dynamics of tritrophic plant–herbivore–parasitoid systems and has important implications for management of herbivorous insect pests.
2. A model system was used to explore the outcome of interspecific competition between parasitoids that differ in host specificity. The system included the lepidopteran pest Heliothis virescens , the generalist parasitoid Cotesia marginiventris , and two specialist parasitoids, Microplitis croceipes and Cardiochiles nigriceps .
3. The generalist, C. marginiventris , dominated intrinsic competition when given an 8-h developmental head start over C. nigriceps or when its oviposition was simultaneous with that of M. croceipes . Microplitis croceipes and especially C. nigriceps larvae prevailed when they were allowed to oviposit prior to C. marginiventris .
4. Rates of host mortality prior to parasitoid emergence varied with parasitoid species composition and with the order of oviposition.
5. Implications for integrated pest management and the adaptive significance of competition as related to host specialisation are discussed.     

Comparative study on the behavioral response to starvation in three species of antlion larvae (Neuroptera: Myrmeleontidae)

To clarify how pit-building antlion larvae behave during prolonged periods of low resource abundance, pit relocation rate, giving-up time, and respiration rate under starvation conditions were examined, using three species of antlion larvae. Most larvae ofMyrmeleon bore never relocated their pits before they starved to death, while larvae ofHagenomyia micans relocated more often thanMyrmeleon formicarius (average number of pit relocations 0.04 forM. bore, 0.19 forM. formicarius, and 0.62/individual/10 days forH. micans). The relative respiration rate, a ratio of respiration rate at starvation to that at satiation, was lower inM. bore andM. formicarius than in H. micans. Thus, there was an inversely proportional relationship between the pit relocation rate and the decrease in respiration rate under starvation conditions in the three species of antlion larvae.     

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.     

Thermal ecology of western tent caterpillars Malacosoma californicum pluviale and infection by nucleopolyhedrovirus

Abstract 1. Western tent caterpillars hatch in the early spring when temperatures are cool and variable. They compensate for sub-optimal air temperatures by basking in the sun.
2. Tent caterpillars have cyclic population dynamics and infection by nucleopolyhedrovirus (NPV) often occurs in populations at high density.
3. To determine whether climatic variation might influence viral infection, the environmental determinants of larval body temperature and the effects of temperature on growth and development rates and larval susceptibility to NPV were examined.
4. In the field, larval body temperature was determined by ambient temperature, irradiance, and larval stage. The relationship between larval body temperature and ambient temperature was curvilinear, a property consistent with, but not necessarily limited to, behaviourally thermoregulating organisms.
5. Larvae were reared at seven temperatures between 18 and 36 °C. Larval growth and development increased linearly with temperature to 30 °C, increased at a lower rate to 33 °C, then decreased to 36 °C. Pupal weights were highest for larvae reared between 27 and 30 °C.
6. The pathogenicity (LD₅₀) of NPV was not influenced by temperature, but the time to death of infected larvae declined asymptotically as temperature increased.
7. Taking into account larval growth, the theoretical yield of the virus increased significantly between 18 and 21 °C then decreased slightly as temperatures increased to 36 °C.
8. Control and infected larvae showed no difference in temperature preference on a thermal gradient. The modes of temperature preference were similar to those for optimal growth and asymptotic body temperatures measured in the field on sunny days.
9. Warmer temperatures attained by basking may increase the number of infection cycles in sunny springs but do not protect larvae from viral infection.     

A plant surface mutation mediates predator interference among ladybird larvae

Abstract.  1. We investigated mechanisms causing predator–predator interference between fourth instar Hippodamia convergens larvae foraging for pea aphids on pea plants, Pisum sativum , with a wild-type wax bloom, and the lack of such interference between larvae foraging on pea plants with a reduced-wax bloom caused by the single gene mutation wel .
2. Observations showed that behavioural interactions between larvae were not affected by wax phenotype. Specifically, larvae did not encounter one another more frequently on normal-wax peas as may have been predicted because reduced ability by coccinellids to attach to normal-wax plant surfaces could restrict them to foraging on only some parts of these plants.
3. In a controlled bioassay on normal-wax peas, H. convergens larvae avoided leaflets previously exposed to another larva. On reduced-wax peas, this effect was not detected.
4. In microcosm experiments, inter-predator interference in terms of prey consumption occurred on normal-wax peas, but not on reduced-wax peas. The interference on normal-wax peas occurred whether two H. convergens larvae were placed on a pea aphid-infested, normal-wax plant simultaneously or sequentially.
5. We conclude that the observed inter-predator interference is not as a result of direct physical contact, but rather arises because of (i) inhibition of foraging by chemical trails left by other larvae, (ii) the inability of larvae to access portions of the normal-wax plants creating aphid refugia, or (iii) a combination of these factors.     

Microhabitat selection by ant-lion larvae Myrmeleon brasiliensis (Návas) (Neuroptera: Myrmeleontidae), in a Forest Reserve, Aquidauana, State of Mato Grosso do Sul

The relative abundance, density, distribution pattern and relation among pit diameter and larvae body size of Myrmeleon brasiliensis Návas were evaluated in two microhabitats: sheltered and exposed. The total of 282 pits were found in sheltered microhabitat and only 50 in the exposed. The density of M. brasiliensis was between one and 43 individuals per m(2). The distribution pattern of larvae tended from pooled to regular distribution as the density increased. In both microhabitats the larvae body size was positively correlated with pit diameter.     

Interference competition in ant-lion (Macroleon quinquemaculatus) larvae

Abstract.

• 1 The effects of density, feeding regime, and body size on interference competition in the pit-digging larvae of the ant-lion Macroleon quinquemaculatus (Hagen) were investigated in laboratory experiments.
• 2 Competition had little effect on the pit size of winners but losers constructed much smaller pits than isolated larvae. Losers were less likely to dig or maintain pits and more likely to move than winners.
• 3 Competition was much stronger between well-fed larvae than between hungry ones, and well-fed competitors showed reduced growth rates. Well-fed larvae orientated themselves so that they could throw sand into their neighbour's pit whereas hungry larvae faced away from each other. Differences in hunger level reversed the competitive advantage of larger larvae only when individuals were of similar size.
• 4 Cannibalism was density-dependent and most frequent in hungry, similar-sized, larvae; the smaller larva was usually the victim.
• 5 Displays/challenges between larvae affected the distance between pits. Body size was the main determinant of contest outcome though pit ownership and hunger level also had an effect.

     

Neither the devil nor the deep blue sea: larval mortality factors fail to explain the abundance and distribution of Tischeria ekebladella

Abstract.  1. The distribution, abundance and population dynamics of herbivorous insects may be affected by trophic interactions, by abiotic influences, or by intra-specific processes. Relatively little is known about how trophic influences vary across space. Here, we investigate spatial variation in mortality in the oak-feeding leaf miner Tischeria ekebladella as attributable to individual causal agents.
2. Leaf miners were experimentally introduced on 67 trees on an island 5 km² in area in south-western Finland. On each tree, some larvae were protected by a muslin bag, others by a glue barrier around the branch and some left exposed.
3. In the bagged transplants, 78.4% of larvae survived, compared with only 9.6% in the other two treatments. Most of the mortality was because of airborne agents: mortality on branches sheltered by a glue barrier was as high as on fully exposed branch tips.
4. We consider mortality caused by parasitoid wasps to be the main source of larval death and the primary factor driving general patterns of survival. The effects of bird predation and premature leaf abscission were negligible.
5. We detected spatial aggregation in larval survival and parasitism rates at the level of individual trees, but not across the landscape.
6. Spatial variation in overall leaf miner survival, parasitism and leaf abscission does not suffice to explain patterns of incidence and abundance of wild T. ekebladella on experimental trees. Rather, we identify metapopulation dynamics as a likely determinant of the spatial distribution of T. ekebladella in the landscape.     

Evolutionarily Stable Relocation Strategy in an Antlion Larva

Antlion pits are often spatially aggregated even though there are potential negative effects associated with the aggregation (e.g., heightened competition and predation risk). This study investigated the possibility that a strategy leading to aggregation can be an evolutionarily stable strategy (ESS). In particular, the strategy considered was ‘decreasing relocation tendency when there are neighbors’. An individual based model showed that the strategy can be the unique ESS when the spatial distribution of prey is not completely random and antlions can learn it from their past foraging experiences. A laboratory experiment was conducted to examine the effects of the presence of neighbors and foraging success on the relocation behavior of antlion larvae. Antlions reduced their relocation tendency with respect to these factors, consistent with the predicted ESS. The results suggest that pit aggregations are formed because antlions reduce their relocation tendency when neighbors exist, and this strategy is an ESS.     

Effects of host-plant shift on immune and other key life-history traits of an eruptive Geometrid, Epirrita autumnata (Borkhausen)

Abstract.  1. Population density of Epirrita autumnata (Lepidoptera: Geometridae) reaches outbreak densities regularly in northernmost Scandinavia. During these outbreak years, the most abundant host species, the mountain birch ( Betula pubescens ssp. czerepanovii ), is regularly exhausted, although larvae may rescue themselves from starvation by using alternative host species.
2. In this paper, the effects of the shift of host species on the immune defence and other life-history traits of E. autumnata were investigated, and possible consequences for population dynamics were briefly discussed. Moth larvae were reared on the leaves of the main host, mountain birch, until larvae reached their third instar. After this, larvae were allocated randomly to five treatments: larvae were either allowed to finish larval stage on the mountain birch or were shifted onto four alternative host species that are typical species for the area.
3. As expected, the host species had a major effect on fitness traits: body weight, development, and survival rate of the moths. The pupal weight was lower and development rates slower on the three alternative host species, Salix myrsinifolia Salisb., Vaccinium uliginosum L., and Betula nana L., than on the main host, mountain birch.
4. The immunity was, however, the same or better on the alternative hosts than on the main host. The immunity and pupal weights were negatively related, suggesting a trade-off between body size and immunocompetence.
5. The decreased body size and fecundity of E. autumnata during outbreak years may be partly due to the shift to alternative host species whereas the host-plant species probably does not affect markedly the rate of parasitism.     

Resistance of kale varieties to attack by Mamestra brassicae

1 The objectives of this work were to study the resistance of six kale ( Brassica oleracea acephala group) varieties to cabbage moth Mamestra brassicae (L.) expressed as antibiosis and to determine the effect of plant age on larval survival and development.
2 The influence of plant age on resistance was determined using leaves from seedlings and from mature plants. Survival and development of M. brassicae larvae and feeding rates were determined in laboratory bioassays.
3 Leaves from seedlings were more suitable than those of mature plants for establishing differences in resistance. There were significant differences between kale varieties in larval survival, growth rate, leaf feeding, and time to pupation but not pupal weight. The varieties MBG-BRS0031, MBG-BRS0351, and MBG-BRS0287 reduced survival of M. brassicae larvae. Larvae that fed on MBG-BRS0060 were the heaviest and took the longest time to pupation. MBG-BRS0031 was consumed significantly less by larvae than were all the other varieties examined. Leaves from mature plants of MBG-BRS0142 and MBG-BRS0170 were defoliated significantly less than those of other varieties.
4 In conclusion, the variety MBG-BRS0031 may be a promising source of resistance to M. brassicae . Leaf antibiotic resistance was shown to play a role in defense against M. brassicae attack but it is not the only possible mechanism of resistance.     

A review of relationships between interspecific competition and invasions in fruit flies (Diptera: Tephritidae)

Abstract.  1. A number of invasions in the family Tephritidae (fruit flies) have been observed worldwide despite quarantine procedures. In this review, the potential importance of interspecific competition and competitive displacement among different tephritid species is evaluated in the context of recent invasions.
2. Where polyphagous tephritid species have been introduced in areas already occupied by a polyphagous tephritid, interspecific competition has resulted in a decrease in number and niche shift of the pre-established species. No reciprocal invasions have been observed.
3. The data on tephritid invasions seem to support a hierarchical mode of competition; however, complete exclusion usually did not occur. Indeed, tephritid distribution and abundance are markedly structured by various abiotic (mostly climatic) and biotic (host plants) factors.
4. The primary determinant of competitive interactions in near-optimal conditions, such as lowlands with abundant fruit plantations, is probably the life-history strategy. The r – K gradient could be used as a predictor of potential invaders, because K traits (such as large adult size) may favour both exploitation and interference competition.
5. For future research, a better understanding of competition mechanisms seems essential. Different species competing in the same area should be compared with respect to: (i) demographic parameters, (ii) the outcome of experimental co-infestations on the same fruit, and (iii) behavioural and chemical interference mechanisms.