Non-interactive multiple predator effects on tadpole survival

Interactions among and within three species of predators were estimated in terms of their effects on prey survival using short-term predation experiments. The prey were tadpoles (Rana temporaria), and the predators were dragonfly larvae (Anax imperator), newts (Triturus alpestris), and backswimmers (Notonecta glauca). Mortality rate per predator imposed by Triturus and Notonecta did not decline with predator density, whereas the predation rate of Anax was strongly reduced when the number of predator individuals increased. Impacts of all three predators were not altered by the presence of other species in pairwise combinations. This system is therefore characterized by interference between individual dragonflies but relatively independent effects of predator species. These results were largely predictable based on the natural history of the predators and are encouraging for attempts to model communities as assemblages of interacting species.     

Species richness along multiple gradients: testing a general multivariate model in oak savannas

Behavioral responses of prey to their predators can critically alter community dynamics. Whether or not a prey responds, clearly depends on the effectiveness of that response. The effectiveness on the other hand is predicted to depend on predator behavior. Actively searching predators can render the behavioral responses in their prey ineffective. Nevertheless, most studies investigating the optimal reaction of prey treated predators as immobile elements of the environment. I experimentally manipulated activity of poolfrog ( Rana lessonae ) tadpoles by keeping them at low and high food levels, and exposed them to three species of invertebrate predators ( Aeshna cyanea, Anax imperator , and Dytiscus marginalis ), whose activity also was manipulated through different food levels. Satiated, less active predators were more likely to kill hungry, more active tadpoles, but hungry predators killed hungry and satiated tadpoles about equally often. This result suggests that reducing their activity is a more effective strategy for tadpoles if the predators themselves are less active. On the other hand, against hungry, highly motivated predators, the behavioral avoidance strategies were essentially ineffective. Antipredator behavior is generally thought to stabilize the dynamics of predator–prey systems. The results presented here, however, suggest that the community dynamical consequences of antipredator behavior also critically depend on decisions made by predators.     

Predation on amphibian eggs and tadpoles by common predators in acidified lakes

Eggs and three different stages of premetamorphic tadpoles of moor frog Rana arvalis and common toad Bufo bufo were offered to the following potential predators in acidified lakes: newt Triturus vulgaris , water beetles Rhantus exoletus and Dytiscus lapponicus , dragonfly larvae Leucorrhinia dubia and Aeshna spp., water boatman Notonecta glauca and water bugs, Cymatia bonsdorffi, Glaenocorisa p. propinqua , and Corixa dentipes.
The predation pressure on eggs of R. arvalis was low due to thick jelly. The eggs of B. bufo were not attractive to predators with chewing mouth parts due to unpalatability but predators with sucking mouth parts were not repulsed. Tadpoles of R. arvalis were eaten by all predators but tadpoles of B. bufo were unpalatable to most predators. The predators used in the experiments are the new top predators in acidified fishless lakes. They may contribute to the reduction of populations of R. arvalis in acidified areas.     

Predation on newt eggs (Triturus alpestris and T. helveticus): identification of predators and protective role of oviposition behaviour

Predation on newt eggs ( Triturus helveticus and T. alpestris ) was studied in both natural (pond) and control (aquaria) conditions. Eggs were laid on artificial supports and set in a pond near Bourg-en-Bresse (south-east France). Survival from segmentation to pre-hatching stages was similar in these two species and estimated at 0·16 ± 0·07. Survival of eggs protected in a closed bag of plastic netting was significantly higher (0·79 ± 0·08), suggesting predation as the major mortality risk. Eggs of T. alpestris were offered to the following potential predators: newts Triturus alpestris and T. helveticus (males and females), adult aquatic insects ( Dytiscus marginalis, Acilius suleatus. Notonecta glauca, Ranatra linearis. Ilyocoris hermanni). a snail ( Lunnaea stugnalis) and a tadpole ( Rana temporaria ). Only the newts and water beetles were observed to cat the eggs. Females wrapped their eggs in a fold made with the support during oviposition. D. marginalis fed on wrapped eggs and those we unwrapped in the same proportion. but newts (especially females) and A. sulcatus clearly ate fewer wrapped than unwrapped eggs. These results are discussed in terms of mortality risk and antipredator adaptations.     

Skeletomuscular development of genital segments in the dragonfly Anax imperator (Odonata, Aeshnidae) during metamorphosis and its implications for the evolutionary morphology of the insect ovipositor

The skeleton-muscular organisation of abdominal segments 7-9 in female Anax imperator L. (Anisoptera, Aeshnidae) was examined in the stages of ultimate larva, teneral imago, and mature imago, with special emphasis on the transformation of the muscle arrangement. The absence of certain muscles in the genital segments compared to the 7th pre-genital segment was noted on all studied stages. Reductions of certain muscles in adults compared to those in larvae are reported. Some of ovipositor's muscles appear already in larvae. Attachment sites of larval muscles are retained in freshly emerged females concurrently with integument transformations. This situation allows for precise determination of the borders of newly differentiated genital sclerites and, therefore, of the possible origin of certain ovipositor elements in odonates. All changes in the segmental sets of studied abdominal muscles during metamorphosis are tabulated, and displacements of muscles are documented and illustrated. Schematic figures illustrating homologies between the parts of larval and imaginal abdominal sclerites are provided. The origins of the components of the endophytic ovipositor in Odonata as well as their implications for the evolutionary morphology of the insect ovipositor are discussed.     

The adipokinetic hormones of Odonata: a phylogenetic approach

Adipokinetic neuropeptides from the corpora cardiaca of the major families of all three suborders of the Odonata were identified by one or more of the following methods: (1) Isolation of the peptides from a methanolic extract of the corpora cardiaca by liquid chromatography, peak monitoring by fluorescence of the Trp residue and comparison of the retention time with those of known synthetic peptides of Odonata. (2) Hyperlipaemic bioassays of the HPLC-generated fractions either in Locusta migratoria or, in a few cases, in Anax imperator or Orthetrum julia. (3) Sequencing of the isolated, bioactive HPLAC fraction by Edman degradation. (4) Mass spectrometric measurement of the isolated, bioactive fraction. Sequence assignment revealed that the investigated Odonata species always contain only one adipokinetic peptide. This is always an octapeptide. The suborder Zygoptera contains the peptide code-named Psein-AKH, the Anisozygoptera and the families Aeshnidae, Cordulegastridae and Macromiidae of the Anisoptera contain Anaim-AKH, whereas Gomphidae, Corduliidae (with the exception of Syncordulia gracilis) and Libellulidae contain Libau-AKH; one species of Libellulidae has Erysi-AKH, a very conservative modification of Libau-AKH (one point mutation). When these structural data are interpreted in conjunction with existing phylogenies of Odonata, they support the following: (1) Zygoptera are monophyletic and not paraphyletic. (2) Anisozygoptera and Anisoptera are sister groups and contain the ancestral Anaim-AKH which is independently and convergently mutated to Libau-AKH in Gomphidae and Libellulidae. (3) The Corduliidae are of special interest. Only Corduliidae sensu stricto appear to contain Libau-AKH, other species placed into this family by most authorities contain the ancestral Anaim-AKH. Possibly, assignments of AKHs can untangle the paraphyly of this family.     

Foraging behaviour influences the outcome of predator–predator interactions

Abstract.  1. Interactions among predators may influence the total efficiency of a predator complex. The effect of intra- and interspecific interactions of the generalist predators Orthotylus marginalis (Heteroptera: Miridae) and Anthocoris nemorum (Heteroptera: Anthocoridae) was investigated in a laboratory experiment. Outcomes of the interactions were determined by comparing predation rates on eggs and larvae of the blue willow beetle Phratora vulgatissima of single individuals with those of two individuals of the same or different species.
2. A non-additive, antagonistic effect on predation rates due to intraspecific interactions was found between individuals of A. nemorum . No such effect was found in O. marginalis . These results are as expected as a consequence of differences in behaviour of the two predator species: A. nemorum is a much more active and mobile predator than O. marginalis .
3. Contrary to expectation, interspecific interactions between A. nemorum and O. marginalis did not affect the total predation rate.
4. An observation from the field corroborated the results obtained in the laboratory study; there was no negative relationship between the densities of the two predator species, indicating that the two species do not interact negatively in the field at their natural densities.
5. It is concluded that the additive effect of multiple predator species is of potential value in biological control.     

Individual vulnerability to predation: the effect of alternative prey types

SUMMARY. 1. Many examples of predator-prey interactions from freshwaters suggest that apparent predator preferences result more from the relative vulnerabilities of prey than any positive choice by the predator.
2. The rank order of vulnerability of seven invertebrate prey to nine invertebrate predators was measured in the laboratory. The ranks were used as a measure of relative vulnerability.
3. The vulnerability of individuals of Simocephalus vetulus (Muller) (Crustacea: Cladocera) to predation by Enallagma cyathigerum (Char-pentier) (Insecta: Odonata) and Notonecta glauca L, (Insecta: Hemiptera) was investigated in the presence of alternative prey of differing relative vulnerabilities, based on the rank order measures.
4. With E. cyathigerum , vulnerability of individual S. vetulus depended greatly on the relative vulnerability of other prey types available. With N. glauca the intrinsic defences of S. vetulus were very effective.
5. The importance of relative prey vulnerability is discussed with particular reference to small habitats with marked annual species turnover.     

Substrate usage and its regulation during flight and swimming in the backswimmer, Notonecta glauca

Abstract.  The metabolites that are generally used by insects during exercise are present in quite different concentrations in the haemolymph of the backswimmer Notonecta glauca L. Lipids are most abundant (between 10 and 20 mg/mL), whereas carbohydrates (2–3 mg/mL) and proline (approximately 1 mg/mL) are at very low concentrations. Injection of an extract of conspecific corpora cardiaca causes pronounced hyperlipaemia in the backswimmer. A neuropeptide with the same effect was isolated from the corpora cardiaca in a single high-performance liquid chromatography (HPLC) step; the primary sequence was deduced from mass spectrometric measurements (matrix-assisted laser desorption/ionization-time of flight and electrospray quadrupol time-of-flight mass spectrometry) of whole corpora cardiaca, and the mass was confirmed in the HPLC fraction that had adipokinetic activity. The biologically active octapeptide has the sequence pGlu-Val-Asn-Phe-Ser-Pro-Ser-Trp amide, which was characterized previously from the corpora cardiaca of the Emperor dragonfly, Anax imperator , and denoted Anaim-adipokinetic hormone (AKH). The synthetic Anaim-AKH peptide causes lipid mobilization when injected at a dose of 1 pmol into N. glauca . When other synthetic AKH members that occur in Hemiptera are injected into N. glauca at the same dose, the hyperlipaemic responses are significantly lower than after injection of Anaim-AKH. Because only lipids increase upon activity, such as continuous swimming for 1 h or during a 1-h rest period after a 3-min flight episode in the laboratory, it is assumed that Anaim-AKH serves as a true adipokinetic hormone in the backswimmer during bouts of natural swimming and flight.     

Changes in feeding behaviour of the larvae of the damselfly Enallagma cyathigerum in response to stimuli from predators

1. The feeding rates, diet composition and diel periodicity in feeding activity among larvae of the damselfly Enallagma cyathigerum Charp., exposed to chemical, visual and mechanical stimuli from the predators Ranatra linearis (L.), Notonecta glauca L. and Rutilus rutilus (L.) were investigated.
2. In response to chemical cues produced by all the predator species, larvae reduced their rate of feeding significantly (especially on large, sedentary prey).
3. Small larvae reacted more strongly than large ones to the kairomone produced by N. glauca.
4. The fish kairomone induced a reduction in feeding activity during daylight hours only. This reaction was more intense than the reaction to non-chemical cues.
5. Observed predator-induced changes in diet composition, caused directly by reduced activity during feeding, are discussed as an antipredator behaviour of damselfly larvae.     

Coding characters from different life stages for phylogenetic reconstruction: a case study on dragonfly adults and larvae,including a description of the larval head anatomy of Epiophlebia superstes (Odonata: Epiophlebiidae)

The exclusive use of characters coding for specific life stages may bias tree reconstruction. If characters from several life stages are coded, the type of coding becomes important. Here, we simulate the influence on tree reconstruction of morphological characters of Odonata larvae incorporated into a data matrix based on the adult body under different coding schemes. For testing purposes, our analysis is focused on a well‐supported hypothesis: the relationships of the suborders Zygoptera, ‘Anisozygoptera’, and Anisoptera. We studied the cephalic morphology of Epiophlebia, a key taxon among Odonata, and compared it with representatives of Zygoptera and Anisoptera in order to complement the data matrix. Odonate larvae are characterized by a peculiar morphology, such as the specific head form, mouthpart configuration, ridge configuration, cephalic musculature, and leg and gill morphology. Four coding strategies were used to incorporate the larval data: artificial coding (AC), treating larvae as independent terminal taxa; non‐multistate coding (NMC), preferring the adult life stage; multistate coding (MC); and coding larval and adult characters separately (SC) within the same taxon. As expected, larvae are ‘monophyletic’ in the AC strategy, but with anisopteran and zygopteran larvae as sister groups. Excluding larvae in the NMC approach leads to strong support for both monophyletic Odonata and Epiprocta, whereas MC erodes phylogenetic signal completely. This is an obvious result of the larval morphology leading to many multistate characters. SC results in the strongest support for Odonata, and Epiprocta receives the same support as with NMC. Our results show the deleterious effects of larval morphology on tree reconstruction when multistate coding is applied. Coding larval characters separately is still the best approach in a phylogenetic framework. © 2015 The Linnean Society of London     

The larval diet of three anisopteran (Odonata) species

SUMMARY.

• 1 Comparisons between larval diets of three anisopteran species (Anax imperator, Aeshna cyanea and Libellula depressa) showed that their food intake changed depending on: (1) predator species, (2) time of year, and (3) developmental stage.
• 2 Although this last factor is not so important, the mean size of prey items and the range of prey species eaten by Auax imperator and Aeshna cyanea larvae increased with predator size.
• 3 Comparisons between prey availability and diets indicated differential selectivity by these predators.

     

Interactions among coexisting larval Odonata: an in situ experiment using small enclosures

Field experiments using small replicated enclosures focused on interactions between larval populations of Epitheca cynosura and Ladona deplanata (Odonata: Anisoptera) — two species that emerge in early spring. The presence of Epitheca reduced the total biomass of Ladona, but Ladona had no significant effect on Epitheca. These early-emerging species reduced the biomass of small instars of late-emerging Anisoptera which colonized enclosures during the experiments; and the late-emerging Anisoptera seem to have inhibited colonization by Zygoptera larvae. Results are consistent with the importance of predatory (cannibalism or mutual predation) interactions in this community.     

Behavioural and life history responses to predation risk by common frog tadpoles exposed to two predators during ontogeny

The presence of predators can induce changes in both the morphology and behaviour of Anuran larvae, affecting both their size and developmental stage at metamorphosis and, consequently, the fitness of adult individuals. Tadpoles have been shown to be capable of finely tuning their defensive responses according to the actual risk perceived, which is expected to vary according to the prey-to-predator size ratio. In this study, we exposed common frog (Rana temporaria) tadpoles (Gosner stages 28–30), for a period of 2 weeks, to the non-lethal presence of dragonfly larvae (Anax imperator) and backswimmers (Notonecta glauca). In such a narrow window of time, we expected behavioural responses to be similar for both predators and exposure to predation risk to have negligible effects on tadpole development and weight. Overall, tadpoles increased hiding behaviour and were less active when predators were present in the experimental mesocosms, but behavioural responses were constrained to the early phase of the ontogeny and were no longer used when tadpoles reached a threshold size. Developmental rate slightly slowed down for predator treatments in comparison to controls, possibly as a consequence of energetic investment in unrecorded morphological defences. Although variation in laboratory conditions and protocols makes it hard to compare the results of different experiments, our results contribute to verify the consistency of behavioural responses in Anuran larvae.     

Predation risk perception and food scarcity induce alterations of life-cycle traits of the mosquito Culex pipiens

Abstract.  1. Predators may affect prey populations by direct consumption, and by inducing defensive reactions of prey to the predation risk. Food scarcity frequently has effects on the inducible defences of prey, but no consistent pattern of food–predation risk interaction is known.
2. In this study the combined effect of food shortage and predation-risk perception in larvae of the mosquito Culex pipiens was investigated. Water exposed to the aquatic predator bug Notonecta glauca was used as a source of predation intimidation. Mosquito larvae were reared in three different media containing either no predator cues or the cues of N. glauca that had been fed on either C. pipiens larvae or on Daphnia magna . Food was provided in favourable or limited amount for these set-ups.
3. The results showed that chemical cues from the predators fed with prey's conspecifics caused a decreased survival, delayed pre-imaginal development, and reduction in body size of emerged mosquitoes, whereas chemical cues from predators fed with D. magna caused only delayed development. Food scarcity significantly exacerbates the negative effect of the predator cues on pre-imaginal development of C. pipiens . Effects of the cues on larval development and body size of imagoes are significantly stronger for females than for males.
4. The present study suggests that when food is limited, predators can affect population dynamics of prey not only by direct predation, but also by inducing lethal and sublethal effects due to perception of risk imposed by chemical cues. To understand the effects of predators on mosquito population dynamics, environmental parameters such as food deficiency should be considered.     

The effect of invertebrate and vertebrate predators on the foraging movements of Ischnura elegans larvae (Odonata: Zygoptera)

SUMMARY.

• 1 The foraging movements of late instar Ischnura elegans larvae were monitored in laboratory experiments to study the effects of predators on larval feeding behaviour.
• 2 Ischnura larvae are sit-atid-wait, or ambush, foragers, moving occasionally between perches in search of profitable feeding sites. Larval foraging movements, monitored at different densities of Daphnia prey, increased significantly when prey were absent.
• 3 In experiments without prey, larval movement was inhibited by the presence of fish predators, as well as by invertebrate predators (Notonecta glauca), but not by closely related, non-predatory invertebrates (Corixa punctata) or physical disturbance of the water (intermittent air bubbles).
• 4 Further experiments varied Ischnura hunger levels (0–8 days without food) and illumination (light or dark) with and without notonectid predators. Hunger had no consistent effect on penultimate instar behaviour but final instar foraging activity was significantly modified: movements increased after 4 days starvation and decreased again after 8 days. This response was suppressed by the presence of predators. Both larval instars moved significantly less often in the light, even when predators were absent.
• 5 These phenotypically flexible predator-avoidance responses are likely to decrease the risk of predation by both visual and tactile predators. However, predators clearly have an important influence on the feeding niche of Ischnura larvae, and may decrease the overall feeding efficiency, growth rate, and survival of larvae by constraining their movement in search of profitable feeding sites.

     

Chemical cues from multiple predator-prey interactions induce changes in behavior and growth of anuran larvae

Chemical signals are used as information by prey to assess predation risk in their environment. To evaluate the effects of multiple predators on prey growth, mediated by a change in prey activity, I exposed small and large bullfrog (Rana catesbeiana) larvae (tadpoles) to chemical cues from different combinations of bluegill sunfish (Lepomis macrochirus) and larval dragonfly (Anax junius) predators. Water was regularly transferred from predation trials (outdoor experiment) to aquaria (indoor experiment) in which activity and growth of tadpoles was measured. The highest predation mortality of small bullfrog larvae in the outdoor experiment was due to Anax, and it was slightly lower in the presence of both predators, probably resulting from interactions between predators. There was almost no mortality of prey with bluegill. The activity and growth of small bullfrog larvae was highest in the absence of predators and lowest in the presence of Anax. In the presence of bluegill only, or with both predators, the activity and growth of small bullfrog tadpoles was intermediate. Predators did not affect large tadpole activity and growth. Regressing mortality of small bullfrog tadpoles against activity and growth of bullfrog tadpoles revealed a significant effect for small bullfrog larvae but a non-significant effect for large bullfrog larvae. This shows that the response of bullfrog tadpoles to predators is related to their own body size. The experiment demonstrates that chemical cues are released both as predator odor and as alarm substances and both have the potential to strongly alter the activity and growth of prey. Different mechanisms by which chemical cues may be transmitted to species interactions in the food web are discussed. Received: 28 June 1999 / Accepted: 15 November 1999     

How Does Environmental Variation Affect the Distribution of Dragonfly Larvae (Odonata) in the Amazon-Cerrado Transition Zone in Central Brazil?

This study investigated the effects of environmental variation on assemblages of dragonfly larvae (Odonata). We hypothesize that there is a significant correlation between species richness, species composition, and abundance of Odonata individuals, and habitat integrity and abiotic variables. To test this hypothesis, we sampled odonate larvae at 12 streams in the Suiá-Miçú River basin in Mato Grosso, Brazil, during three different periods of the year. Local physical and chemical variables (temperature, pH, turbidity, electrical conductivity (EC), dissolved oxygen (DO), total dissolved solids (TDS), and oxidation reduction potential (ORP)) were measured at each site using a multi-parameter probe, and integrity was assessed using the Habitat Integrity Index (HII). The variation in richness, abundance, and composition of Odonata species was related to the environmental variables analyzed, primarily by the abiotic factors pH, electrical conductivity, dissolved oxygen, total dissolved solids, and oxidation reduction potential. Our hypothesis was corroborated for the suborder Anisoptera, which showed a significant relationship with these variables, whereas Zygoptera was only related to pH. Our results show the importance of physical and chemical conditions in ecological studies using Odonata larvae as tools for the management and conservation of freshwater ecosystems.     

Lake acidity,fish predation,and the distribution and abundance of some littoral insects

Densities of Corixidae (Hemiptera), larval Odonata, and large larval Trichoptera were estimated in the littoral zone of small lakes in an acid-stressed area near Sudbury, Ontario. Fish were present in some lakes and absent in others, and fishless lakes occurred across a wide range of pH. Corixidae were significantly more abundant in lakes without fish than in lakes with fish, and their numbers were not related to the pH of fishless lakes. Anisoptera (Odonata) larvae tended to be more numerous in benthic samples from fishless lakes than from lakes with fish, and their exuviae were significantly more abundant around fishless lakes. In most lakes, the assemblage was dominated by three species; Leucorrhinia glacialis, Libellula julia, and Cordulia shurtleffi. In lakes containing white sucker, Catostomus commersoni, Gomphus spp. were most numerous. In the most acid fishless lakes, L. julia was uncommon, and L. glacialis was extremely abundant. In fishless lakes, numbers of Anisoptera larvae and exuviae were negatively correlated with pH, though species richness was positively correlated with pH. Exuviae of Zygoptera (Odonata) were more abundant around fishless lakes, irrespective of pH. Larvae of Limnephilus (Trichoptera) were most abundant in non-acid fishless lakes, and absent at pH<5.2. Abundances of Banksiola (Trichoptera) were negatively correlated with the pH of fishless lakes.     

Effects of interspecific competition, predation, and their interaction on survival and development time of immature Anopheles quadrimaculatus.

We examined the effect of predation by the backswimmer (Notonecta undulata; Hemiptera: Notonectidae), competition by zooplankton and snails, and both predation and competition on the survival and development time of larval Anopheles quadrimaculatus mosquitoes in experimental mesocosms. We found that both predation and interspecific competition greatly reduced the survivorship of larvae and the number of larvae emerging into adulthood. Treatments with both predators and competitors had fewer larvae transitioning among instars and into adulthood but not in an additive way. In addition, mosquito larvae in the presence of predators and competitors took two days longer to emerge than where predators and competitions were absent. Our work provides evidence that biotic interactions, such as predation and competition, can strongly regulate the number of mosquito larvae by reducing the number of larvae that survive through instars and to emergence and by increasing the generation time.