Antlion Larvae Research Articles

A trade-off between growth and starvation endurance in a pit-building antlion

Trade-offs have a central role in evolutionary ecology and life-history theory. Here, we present evidence for the existence of a rarely studied trade-off between growth rate and starvation endurance in larvae of a pit-building antlion. We first manipulated antlions' feeding regime and obtained a spectrum of growth rates. Next, we starved the antlions and documented their rate of mass loss. Antlions growing faster during the feeding phase also lost mass faster during the successive starvation period, implying the existence of an induced trade-off between fast growth and starvation endurance. Finally, we fed all antlions with prey items of similar mass and measured both the giving-up prey mass (i.e. the remaining body mass of the prey that was not converted into predator body mass), and growth efficiency of antlions (i.e. proportion of prey consumed, negatively correlated with giving-up prey mass). The giving-up mass was negatively correlated with the growth rate of the antlions during the feeding phase, and positively correlated with their growth rate during the starvation phase (the opposite pattern was evident when examining growth efficiency), incongruently with the common phenomenon of growth compensation (i.e. extracting more of the prey after a starvation period). We suggest that antlion larvae can adopt a physiological mode bounded by two extremes: one extreme is adapted to starvation, involving reduced metabolic rates but also reduced capability to exploit prey, while the other is adapted to fast growth, allowing an efficient exploitation of prey, but at the expense of lowered starvation endurance.

Substrate particle size-preference of wormlion Vermileo vermileo (Diptera: Vermileonidae) larvae and their interaction with antlions

Wormlion larvae are found in substrates consisting of fine sand or powder, implying that they may be able to distinguish between different substrates according to particle size. To estimate the effects of particle size on wormlions, the pit-building decision of the larvae of the wormlion Vermileo vermileo was observed in four substrates consisting of different sand fractions. Wormlion larvae prefer the finest sand fraction with particle size <= 230 µm. When wormlions (Vermileo vermileo) and antlions (Euroleon nostras) are placed in the same container with two different substrates, interspecific predation does not occur. In two-substrate choice tests larvae of the two species show opposite preferences for two substrates offered. While wormlion larvae readily build pits in the finest sand fraction (<= 230 µm), antlion larvae prefer coarser sand (with particle size 230-540 µm). Wormlion preference for the finest sands and powders, and antlion preference for sands of medium particle size was confirmed by field observations. Sand particle size affects the spatial distribution of sand-dwelling insect larvae and thus may reduce conflicts between heterospecifics.

Foraging behaviour and habitat selection in pit-building antlion larvae in constant light or dark conditions

Pit-building antlion larvae are small sit-and-wait arthropod predators that dig conical pits in sandy soils. We investigated the effect of exposure to constant light versus constant dark conditions on antlion behaviour. Antlions tended to relocate less often, construct pits more frequently and construct larger pits in constant light. We interpret this as an indication of dichotomous behaviour under light versus dark conditions such that antlion larvae hunt during the day and modify their positions at night. Antlion larvae immediately responded to the complete switch in this illumination treatment and adapted their behaviour to fit the new conditions. Prey capture success did not differ between the constant light and constant dark conditions. When provided with a choice between light and dark conditions, the vast majority of larvae preferred the former. Those that preferred constant dark, however, were on average larger than those that preferred constant light. We thus suggest that since larger antlion larvae possess larger fat reserves, they can afford reductions in foraging activity (possibly to increase safety). An examination of the antlion trade-off between their preference for light and their preference for sand depth (which may affect their ability to evade predators) showed that their response to illumination was much more prominent, irrespective of the sand depth. Finally, combining light with other factors such as temperature and density may further elucidate microhabitat selection in pit-building antlion larvae.

Insecticidal bacterium isolated from an ant lion larva from Munakata, Japan

Twenty bacteria were isolated from four ant lion larvae. The isolates were classified into three groups by biological characteristics. Since Group I, Group II and Group III were isolated from individual larvae Kuo1, Kuo3, 4 and Kuo2, respectively, with exception of one isolate Kuo2-1, each ant lion tested had its own dominant bacterial flora. Groups I and II were closer to Serratia liquefaciens and Enterobacter cloacae, respectively, whereas Group III could not be identified by the test used. The phylogenetic analysis of GroEL amino acid sequences revealed that Group I, II and III were related to those of Serratia spp., E. cloacae and Salmonella spp. –Escherichia/Shigella spp., respectively. Among these groups, Group I was highly virulent against Bombyx mori and Periplaneta americana, and caused 100% mortality within 24 h. The other two groups (Group II and III) were avirulent to these insect species. The culture filtrate of Group I caused killing activity to B. mori larvae and the insecticidal substance was purified from culture filtrate of Group I bacterium. Since the insecticidal activity highly correlated with proteolytic activity in the chromatographies, Group I bacterium may secret insecticidal proteinase in vitro.

Variation in food availability influences prey‐capture method in antlion larvae

Abstract 1. Larvae of a Myrmecaelurus sp. are unique among antlions because they have two prey‐capture methods; they either ambush prey at the surface, or dig pit traps that prey fall in to. It was hypothesised that larvae will use the capture method that maximises their net rate of energy gain, which will be influenced by food availability (encounter rate) and by past energy inputs (body condition).2. Costs were estimated by measuring resting and activity metabolic rates and determining the duration of pit maintenance at various encounter rates with ants that served as prey. Benefits were estimated from the energy gained per ant captured at different encounter rates.3. Net energy gained was higher with a pit than without one, and was influenced more by the differences in prey capture rate between the two capture methods, and less by the differences in energy costs associated with each method. The proportion of larvae that constructed pits was higher when they were in intermediate body condition than when in good or in poor body condition.4. Thus, the use of one capture method or the other depends on a combination of the influences of past net energy gain and the antlion’s most recent change in encounter rate with prey. Ambushing without a pit may serve as a default when physiological constraints limit the larvae’s ability to invest in pit construction and maintenance, or when larvae are sated, and saving the energy of pit construction and maintenance is worthwhile.

Effect of spatial pattern and microhabitat on pit construction and relocation inMyrmeleon hyalinus(Neuroptera: Myrmeleontidae) larvae

Abstract1. In two sets of enclosure experiments, we studied the spatial pattern, relocation rates, pit construction rates, and microhabitat preference ofMyrmeleon hyalinuslarvae.2. We showed thatM. hyalinuslarvae actively prefer shady sites and often relocate to shady areas when exposed to the sun. This behaviour may constitute a life‐saving strategy in desert environments.3. The initial spatial pattern in a cluster of antlion larvae did not affect the final pattern, relocation rate, or the pathway moved while relocating. We interpret this finding to mean that the spatial pattern ofM. hyalinuslarvae is mainly influenced by factors operating at large time‐scales, such as exploitation competition and physical‐microclimatological constraints, rather than those operating at shorter time‐scales, such as interference competition.4. In contrast, the likelihood of pit construction was positively correlated with nearest neighbour distance, possibly as a result of interference competition.5. Pit construction rates were constant throughout the experiment, while relocation rates decreased with time.6. Contrary to previous studies, we found no correlation between body mass and spatial position inside the cluster, suggesting that these differences are the products of slower processes (e.g. exploitation and body condition).

The interplay between foraging mode, habitat structure, and predator presence in antlions

Antlion larvae are sand-dwelling insect predators, which ambush small arthropod prey while buried in the sand. In some species, the larvae construct conical pits and are considered as sit-and-wait predators which seldom relocate while in other species, they ambush prey without a pit but change their ambush site much more frequently (i.e., sit-and-pursue predators). The ability of antlion larvae to evade some of their predators which hunt them on the sand surface is strongly constrained by the degree of sand stabilization or by sand depth. We studied the effect of predator presence, predator type (active predatory beetle vs. sit-and-pursue wolf spider), and sand depth (shallow vs. deep sand) on the behavioral response of the pit building Myrmeleon hyalinus larvae and the sit-and-pursue Lopezus fedtschenkoi larvae. Predator presence had a negative effect on both antlion species activity. The sit-and-wait M. hyalinus larvae showed reduced pit-building activity, whereas the sit-and-pursue L. fedtschenkoi larvae decreased relocation activity. The proportion of relocating M. hyalinus was negatively affected by sand depth, whereas L. fedtschenkoi was negatively affected also by the predator type. Specifically, the proportion of individual L. fedtschenkoi that relocated in deeper sand was lower when facing the active predator rather than the sit-and-pursue predator. The proportion of M. hyalinus which constructed pits decreased in the presence of a predator, but this pattern was stronger when exposed to the active predator. We suggest that these differences between the two antlion species are strongly linked to their distinct foraging modes and to the foraging mode of their predators.

Direct and indirect effects of soil structure on the density of an antlion larva in a tropical dry forest

Abstract 1. We used structural equation models to discriminate direct and indirect effects of soil structure on the abundance of the antlion Myrmeleon crudelis, a neuropteran larva that digs conical pits in soil to capture small arthropods. We proposed that soil structure may modify antlion density indirectly through its influence on tree cover, which in turn directly alters the amount of sun and rain that can reach the forest floor and the amount of litter fall.2. The proportion of finer soils positively affected antlion density directly, but negatively tree cover. Tree cover positively affected both the amount of leaf litter and antlion density. Leaf litter negatively affected antlion density. The indirect effects of soils varied in strength and sign depending on whether trees are considered shelters against sun and rain, or leaf litter sources. The relative importance of these effects might also vary between years and seasons.3. Antlions may select patches of finer soils not only because they are easy substrates in which to build pits, but also for their indirect benefit as sites with low leaf litter, illustrating how indirect interactions may affect the local abundance of semi‐sedentary insects.

Orientation of the pit-building antlion larva Euroleon (Neuroptera, Myrmeleontidae) to the direction of substrate vibrations caused by prey

Pit-building antlion Euroleon nostras constructs efficient traps in sand to catch its prey. The predator is known to react to substrate vibrations produced by movements of its prey outside the pit with sand-tossing behaviour but it has not yet been ascertained if this reaction is directed towards the prey. The accuracy of the sand-tossing response in the presence of four prey species was measured using a video recording method. The sand-tossing angle was highly positively correlated with the prey angle. Sand tossing was most frequently elicited when prey was on the posterior sand surface. Covering the larval photoreceptors did not influence the antlion's localizing behaviour.

Why are pitfall traps so rare in the natural world?

Prey capture by trapping is uncommon taxonomically, and generally requires highly evolved cognitive powers (humans) or specialist self-secreted materials (for example, spiders and caddisfly larvae). The most notable exception to this is the conical traps dug by antlion larvae. The relative uncommonness (taxonomically and ecologically) of such pitfall traps has been described as an unexplained mystery in recent publications. Here we suggest some potential routes that might lead to resolution to this mystery. We argue that although such pitfall traps have numerous benefits and are relatively cheap and easy to construct, they may suffer two significant disadvantages relative to, for example, spiders’ webs. First, pitfall traps may require a quite specialist microhabitat. Second, antlion pitfall traps may only work to retain all but the smallest prey if the antlion is present at the bottom of the pit. Thus, antlion may be more functionally tied to their trap than spiders and (since traps are much more visually conspicuous than their owners) this may make them vulnerable to predators and parasitoids that cue on the traps. Both these hypothesised drawbacks are speculative in the absence of a strong body of data and so we discuss how both potential costs could be explored empirically.

Orientation towards prey in antlions: efficient use of wave propagation in sand

Substrate-borne vibration for locating mates, predators and prey is widespread in the animal kingdown. Antlion larvae dig funnel-shaped traps to catch ants and they are totally immersed in dry sand. We used a playback setup reproducing an ant walking on sand to clearly demonstrate that antlions use sand-borne vibrations to locate their prey. Half the tested animals moved towards the stimulus source. The shoot angle of sand tossing was very close to the target angle, indicating excellent ability to perceive stimulus direction. We also discuss orientation mechanisms in sand, a medium with highly unusual wave propagation properties.

Consumption of introduced prey by native predators: Argentine ants and pit-building ant lions

When populations of native predators are subsidized by numerically dominant introduced species, the structure of food webs can be greatly altered. Surprisingly little is known, however, about the general factors that influence whether or not native predators consume introduced species. To learn more about this issue, we examined how native pit-building ant lions (Myrmeleon) are affected by Argentine ant (Linepithema humile) invasions in coastal southern California. Compared to areas without L. humile, invaded areas contained few native ant species and were deficient in medium-sized and large bodied native ants. Based on these differences, we predicted that Argentine ants would negatively affect ant lion larvae. Contrary to this expectation, observational surveys and laboratory growth rate experiments revealed that Myrmeleon were heavier, had longer mandibles, and grew more quickly when their main ant prey were Argentine ants rather than native ants. Moreover, a field transplant experiment indicated that growth rates and pupal weights were not statistically different for larval ant lions reared in invaded areas compared to those reared in uninvaded areas. Argentine ants were also highly susceptible to capture by larval Myrmeleon. The species-level traits that presumably make Argentine ant workers susceptible to capture by larval ant lions—small size and high activity levels—appear to be the same characteristics that make them unsuitable prey for vertebrate predators, such as horned lizards. These results underscore the difficulties in predicting whether or not numerically dominant introduced species serve as prey for native predators.

The Energetic Determination, Spatial Dispersion and Density Dependence of Myrmeleon Ant Lion Pits in Las Cruces, Costa Rica

ABSTRACTThe amount of space used by an organism is energetically determined. We utilized a population of ant lion larvae in Costa Rica to test allometric theories concerning the use of space by organisms and how different densities of individuals affect the use of space. The area of ant lion trapping pits scaled with mass to the three‐quarters power, supporting allometric theory for sessile organisms. Our analyses also show that larger ant lion larvae show spatial repulsion and facultative density dependent pit‐building strategies.

Desenvolvimento de Myrmeleon brasiliensis (Navás) (Neuroptera, Myrmeleontidae), em laboratório, com diferentes dietas naturais

Antlions larvae are sit-and-wait predators that capture arthropod prey in conical sand pits. The aim of this paper were to determine the effect of different natural diets [leaf-cutting ants ( Atta spp.), fruit fly larvae ( Anastrepha spp. and Ceratitis capitata ) and mixed diet ( Atta spp. plus fruit fly larvae)] on the development of larvae and pupae of M. brasiliensis (Navas, 1914) and to estimate the number and size of prey caught in each instar and on each kind of diet. The average duration (days) of the second and third instars of M. brasiliensis was longer when larvae of antlion were fed with leaf-cutting ants. The diets did not affect the duration of the pupal stage nor the pupae size. The different diets did not affect the mean width of head capsule (mm), the mean weight (mg) or the mean body size (mm) in the different instars of M . brasiliensis . In the second and third instars, the larvae of M. brasiliensis fed with leaf-cutting ants consumed more prey than larvae kept on other diets. Adults whose larvae were fed fruit fly larvae were larger than adults on other diets. Although Myrmeleontidae are few studied in Brazil, these results contribute to knowledge of M. brasiliensis biology, but also suggest the need of studies about the development of larvae and pupae in natural environments.

Development of Myrmeleon brasiliensis (Navás) (Neuroptera, Myrmeleontidae), in laboratory, with different natural diets

Antlions larvae are sit-and-wait predators that capture arthropod prey in conical sand pits. The aim of this paper were to determine the effect of different natural diets [leaf-cutting ants (Atta spp.), fruit fly larvae (Anastrepha spp. and Ceratitis capitata) and mixed diet (Atta spp. plus fruit fly larvae)] on the development of larvae and pupae of M. brasiliensis (Navás, 1914) and to estimate the number and size of prey caught in each instar and on each kind of diet. The average duration (days) of the second and third instars of M. brasiliensis was longer when larvae of antlion were fed with leaf-cutting ants. The diets did not affect the duration of the pupal stage nor the pupae size. The different diets did not affect the mean width of head capsule (mm), the mean weight (mg) or the mean body size (mm) in the different instars of M. brasiliensis. In the second and third instars, the larvae of M. brasiliensis fed with leaf-cutting ants consumed more prey than larvae kept on other diets. Adults whose larvae were fed fruit fly larvae were larger than adults on other diets. Although Myrmeleontidae are few studied in Brazil, these results contribute to knowledge of M. brasiliensis biology, but also suggest the need of studies about the development of larvae and pupae in natural environments.

Efficiency of antlion trap construction

Assessing the architectural optimality of animal constructions is in most cases extremely difficult, but is feasible for antlion larvae, which dig simple pits in sand to catch ants. Slope angle, conicity and the distance between the head and the trap bottom, known as off-centring, were measured using a precise scanning device. Complete attack sequences in the same pits were then quantified, with predation cost related to the number of behavioural items before capture. Off-centring leads to a loss of architectural efficiency that is compensated by complex attack behaviour. Off-centring happened in half of the cases and corresponded to post-construction movements. In the absence of off-centring, the trap is perfectly conical and the angle is significantly smaller than the crater angle, a physical constant of sand that defines the steepest possible slope. Antlions produce efficient traps, with slopes steep enough to guide preys to their mouths without any attack, and shallow enough to avoid the likelihood of avalanches typical of crater angles. The reasons for the paucity of simplest and most efficient traps such as theses in the animal kingdom are discussed.

Predatory insects as bioindicators of heavy metal pollution

Heavy metal concentrations of different predatory insects were studied near by a steel factory and from control sites. Waterstriders (Gerridae), dragon fly larvae (Odonata), antlion larvae (Myrmeleontidae) and ants (Formicidae) were analyzed by AAS. In most cases the metal concentrations were higher near the factory, but e.g. waterstriders had higher cadmium concentrations in control area. Discriminant analysis clearly reveals that all these insect groups can be used as heavy metal indicators. However, the commonly used ants were the least effective in indicating the differences between the factory and control sites. Waterstriders are good in detecting differences in iron and manganese, but seem to be poor in accumulating nickel and lead. Antlions are efficient in detecting differences in iron. Antlions and ants are effective in accumulating manganese; as well antlions are efficient in accumulating cadmium. Waterstriders are poor in accumulating lead, but antlions and ants are effective.

Factors Influencing Site Abandonment and Site Selection in a Sit-and-Wait Predator: A Review of Pit-Building Antlion Larvae

There is a large body of evidence indicating that predator behavior may strongly influence patterns and processes at the population and community level. Site selection is a major component of fitness in sit-and-wait predators, especially when relocation is rare. Although several review articles dealt with these issues in web-building spiders, this is the first attempt to summarize the effects of biotic and abiotic factors on site selection and relocation in another group of sit-and-wait predators, the pit-building antlions (Neuroptera: Myrmeleontidae). Our synthesis shows that prey abundance may have relatively little effect on pit relocation and that physical properties of the habitat or competition often override its effect. We suggest that owing to a variety of constraints such as physiological constraints or difficulties in assessing site quality, site selection and relocation are not necessarily optimal and thus food intake rate is not maximized. We call for a multi-factorial study on a single species in order to pinpoint the dominant factors and to assess to what extent they influence site selection and relocation. We conclude by proposing new research directions, such as studying whether pit relocation is an adaptive response, when controlling for possible phylogenetic effects.

Pit building ant-lion larvae effect to the distribution of the substrate particles in their microhabitats

The distribution of the size of the substrate particles in and around the pits of the pit-building antlion larvae was examined at two different types of microhabitats (open and protected). Samples of substrate were taken from the side of the pits, from the edges of the pits and from 3 cm and 10 cm away from the edges of the pit. The size of the substrate particles taken 10 cm away from the edge is considered to be characteristic for the microhabitat. The samplings consisted of different quality of substrates such as sand, loess, clay and they were taken at all three larvae stages. The samples were divided into fractions by a sieve series and the distribution of the weight percentage of the granulation was analyzed. In order to compare the samples the cluster analysis and statistical tests were used. The preference calculations were performed with Ivlev's index of preference (Ei minimum -1; Ei maximum +1). In this respect the distribution of sandy area in the Carpathian Basin is in correspondence with the distribution of the pit-building ant-lion larvae living in open habitats. The preference of the ant-lion larvae living in protected habitat does not depend on the bedrock. The investigation proved that the substrate has particular spatial distribution in the pits and their environment (microhabitat). While building and repairing the pit, the larvae gather the fine particles in the pit and throw the bigger particles further out from the pit. The spatial distribution of the particle size of the substrate can be actively influenced by the chitin bristles and teeth of the mandibles and head of the larvae. Statistical analysis has proved that the particle size preference does not depend on the development stage of the larvae, it is not species dependent and it does not depend on the quality of the substrate as well. The preferred particle size of the pit-building ant-lion larvae was determined by the physical characteristics (particle size, to what extent they were cemented) of the substrate (sand, loess, clay) which is typical of the habitat. The present and the earlier studies suggest that Euroleon nostras, and very likely other pit-building species living in protected microhabitat, chose their habitats providing less food besides optimal particle size and thermal conditions. On the other hand, the species occurring in open habitats such as Myrmeleon bore, live in habitats with bigger food supply but with extreme thermal conditions and coarser particle size.

Substrate particle size affects pit building decision and pit size in the antlion larvae Euroleon nostras (Neuroptera: Myrmeleontidae)

Abstract. The larvae of the antlion Euroleon nostras are pit‐builders, constructing pitfall traps in loose sand. The number of pits and the pit diameter are recorded when larvae are kept in substrates with different particle sizes. The most convenient pit‐building sand fractions are two fractions with fine sand (≤ 0.23 mm; 0.23–0.54 mm). The largest pits are constructed in sand with a particle size of 0.23–0.54 mm. In this sand fraction, larvae of all three instars most readily build pits. No pits are constructed in sand with a particle size greater than 1.54 mm. First‐ and second‐instar larvae avoid building pits in substrates of particle size 1–1.54 mm, but third‐instar larvae construct pits in this sand fraction. It is assumed that the antlion is capable of distinguishing between substrate types and this hypothesis is tested by giving larvae the choice of building a pit in one of four particle‐size fractions. Larvae of all three instars prefer to build pits in the fraction with a particle size of 0.23–0.54 mm. Only third‐instar larvae build pits in all four fractions, but only occasionally in the coarser fraction.