Responses to worm-like-wriggling models by the praying mantis: effects of amount of motion on prey recognition

Adult females of the mantis, Tenodera angustipennis, were presented with a wriggling model, consisting of six circular spots positioned in a row horizontally and adjacently. During presentation, this model wriggled like a worm by moving some spots. When the motion of the model was small (the number of moving spots ≤2), the mantis sometimes stalked the model with peering movements but seldom struck it. When the motion was large (the number of moving spots ≥3), the mantis frequently fixated, rapidly approached, and struck the model. These results suggest that the mantis changes its approach behavior depending on the amount of prey motion. Disappearance of some terminal spots at the stationary end hardly affected the rates of fixation, peering, and strike. The model that wriggled at each end elicited lower rates of fixation and strike than the model that wriggled at one end. These results suggest that the mantis responds to only the fastest moving part of the wriggling model when the motion of the model is large. Electronic Publication     

Predatory behavior of the praying mantis,Tenodera aridifolia I. Effect of prey size on prey recognition

Predatory behavior and effect of prey size of the praying mantis,Tenodera aridifolia, was investigated using three-dimensional prey models. Parameters of prey such as width, length, shape (length/width), area (width×length) and volume were investigated for effects on Watch and Attack responses. Regression analyses suggest that volume is the most important variable in prey recognition. This behavioral trait is considered adaptive, since prey volume is the most reliable measurement for the energy content of the prey.     

The midline metathoracic ear of the praying mantis,Mantis religiosa

Summary The praying mantis, Mantis religiosa, is unique in possessing a single, tympanal auditory organ located in the ventral midline of its body between the metathoracic coxae. The ear is in a deep groove and consists of two tympana facing each other and backed by large air sacs. Neural transduction takes place in a structure at the anterior end of the groove. This tympanal organ contains 32 chordotonal sensilla organized into three groups, two of which are 180° out of line with the one attaching directly to the tympanum. Innervation is provided by Nerve root 7 from the metathoracic ganglion. Cobalt backfills show that the auditory neuropile is a series of finger-like projections terminating ipsilaterally near the midline, primarily near DC III and SMC. The auditory neuropile thus differs from the pattern common to all other insects previously studied.     

Predatory behavior of the praying mantis,Tenodera aridifolia II. Combined effect of prey size and predator size on the prey recognition

Predatory behavior of the praying mantis,Tenodera aridifolia, as a function of the combined effect of its size and the size of the prey was investigated by using prey models. Behavioral responses were almost identical through the nymphal development in the predator. As the mantis grew, it attacked larger prey models, suggesting that it recognizes the prey's size in accordance with its own body size. Regression analyses demonstrate that the ratio of the prey's volume to the cube and the square of the predator's length is a more important parameter for prey recognition than are the one-dimensional parameters of the prey's and the predator's sizes.     

Responses to non-locomotive prey models by the praying mantis,Tenodera angustipennis Saussure

Adult females of the praying mantisTenodera angustipennis were presented with computer-generated images, and the attractiveness of “non-locomotive” prey models was examined. Mantises fixated and struck the “body and leg” model (consisting of an immobile black square on a white background with 2 black lines oscillating randomly at its sides) more frequently than the “leg” model (only oscillating lines) or the “body” model (static square only). This indicates that the model consisting of a static object and moving lines effectively elicits mantis strike behavior, although it is “non-locomotive.”     

Responses of descending neurons to looming stimuli in the praying mantis <Emphasis Type="Italic">Tenodera aridifolia</Emphasis>

Responses to visual stimuli of some neurons that descend the nerve cord from the brain were recorded extracellularly in the mantis Tenodera aridifolia. Most of the recorded neurons showed their largest responses to looming stimuli that simulated a black circle approaching towards the mantis. The neurons showed a transient excitatory response to a gradually darkening or receding circle. The neurons showed sustained excitation to the linearly expanding stimuli, but the spike frequency decreased rapidly. The responses of the neurons were affected by both the diameter and the speed of looming stimuli. Faster or smaller looming stimuli elicited a higher peak frequency. These responses were observed in both recordings from the connective between suboesophageal and prothoracic ganglia and the connective between prothoracic and mesothoracic ganglia. There was a one-to-one correspondence of spike firing between these two recordings with a fixed delay. The neurons had the receptive field on ipsilateral side to its axon at the cervical connective. These results suggest that there is a looming-sensitive descending neuron, with an axon projecting over prothoracic ganglion, in the mantis nervous system.     

Defence behaviours of the praying mantis Tenodera aridifolia in response to looming objects

Defence responses to approaching objects were observed in the mantis Tenodera aridifolia. The mantis showed three kinds of behaviour, fixation, evasion and cryptic reaction. The cryptic reaction consisted of rapid retraction of the forelegs under the prothorax or rapid extending of the forelegs in the forward direction. Obstructing the mantis’ sight decreased its response rates, suggesting that the visual stimuli generated by an approaching object elicited the cryptic reaction. The response rate of the cryptic reactions was highest for objects that approached on a direct collision course. Deviation in a horizontal direction from the direct collision course resulted in a reduced response. The response rate of the cryptic reaction increased as the approaching velocity of the object increased, and the rate decreased as the object ceased its approach at a greater distance from the mantis. These results suggest that the function of the observed cryptic reactions is defence against impending collisions. The possible role of the looming-sensitive neuron in the cryptic reaction is also discussed.     

Coordinated movements of the head and body during orienting behaviour in the praying mantis Tenodera aridifolia

The visual orienting behaviour towards prey in the free-moving mantis was investigated with a high-speed camera. The orienting behaviour consisted of head, prothorax, and abdomen rotations. Coordinated movements of these body parts in the horizontal plane were analysed frame-by-frame. Rotations of these body parts were initiated with no or slight (≤40 ms) differences in timing. The initiation timing of prothorax-abdomen rotation was affected by its initial angle before the onset of orienting. There were positive correlations in amplitude among head-prothorax, prothorax-abdomen, and abdomen rotations. The ratio of these rotations to total gaze rotation was affected by the initial prothorax-abdomen angle before the onset of orienting. Our data suggest that coordinated movements of the head, prothorax, and abdomen during orienting are ballistic events and are pre-determined according to visual and proprioceptive information before the onset of orienting.     

The antennal sensilla of the praying mantis Tenodera aridifolia: A new flagellar partition based on the antennal macro-, micro- and ultrastructures

In insects, the antenna consists of a scapus, a pedicellus, and a flagellum comprising many segments (flagellomeres). These segments possess many morphological types of sensory organs (sensilla) to process multimodal sensory information. We observed the sensilla on flagellomeres in praying mantis (Tenodera aridifolia) with both scanning and transmission electron microscopes. We classified the sensilla into six types: chaetic, campaniform, coelocapitular, basiconic, trichoid and grooved peg sensilla, and inferred their presumptive functions on the basis of their external and internal structures. In addition, based on their distribution, we newly divided the flagellum into 6 distinct parts. This new division leads to a better understanding about the sexual dimorphism and the antennal development in the mantises. The sexual difference in distribution of the grooved peg sensilla suggests that this type of sensilla may play a role in sex-pheromone detection in mantis, which is a rare case of double-walled sensilla mediating this function.     

Visual deprivation and distance estimation in the praying mantis larva

Abstract. Young larvae of the praying mantis, Tenodera sinensis Saussure, were placed on an off-centre island surrounded by a round arena with six black bars painted on a white inner wall. In this situation, it was shown that the horizontal peering movements of the head often seen in mantids are in fact used to measure distances; motion parallax may be involved in this process. Aimed jumps that followed peering were taken to be the distinct result of an absolute distance measurement. Specific visual deprivation such as painting over of certain parts of the eye with opaque black varnish or degeneration of the fovea with sulforhodamine showed that: absolute evaluation of distance is only possible with two fully intact eyes; the peering mechanism is under visual control; and visual experience has a long-term effect on distance measurement involving peering movements.     

Apparent decoupling of prey recognition ability with prey availability in an insular snake population

Numerous studies on the feeding behavior of snakes have reported the consistency of tongue-flick responses with their natural diets. For representatives of widely distributed, dietary generalist species from particular localities, we can expect that their tongue-flick responses to potential prey unavailable in their original habitats have been reduced whereas those to prey common in the habitats have been enhanced. To test this hypothesis, intraspecific variation in tongue-flick responses to prey chemicals was examined using ingestively naive snakes (Elaphe quadrivirgata) from dietarily different populations: populations from the main Japanese island, where the snakes' diet predominantly consists of sympatric frogs, and from Mikura-jima Island, where no frogs occur and the snakes thus chiefly prey on lizards. We presented chemical stimuli from six items including those from their natural and potential prey (fish, frog, lizard, mouse, water, and cologne) to newborn snakes. Significant effects of stimuli on the tongue-flick responses were detected. On the other hand, effects of population and interaction between stimuli and population were not significant, and individual comparisons revealed no significant interlocality differences in responses to either frog or lizard chemicals. Thus, our hypothesis was not supported. However, in the Mikura-jima sample, significantly fewer snakes responded to frog chemicals than in the main island sample. The significance of the inconsistency between prey recognition ability and prey availability in the Mikura-jima population are discussed. Received: October 17, 2000 / Accepted: December 14, 2000     

Response of predators to prey abundance: separating the effects of prey density and patch size

We tested the relative and combined effects of prey density and patch size on the functional response (number of attacks per unit time and duration of attacks) of a predatory reef fish (Cheilodactylus nigripes (Richardson)) to their invertebrate prey. Fish attacked prey at a greater rate and for longer time in large than small patches of prey, but large patches had naturally greater densities of prey. We isolated the effects of patch size and prey density by reducing the density of prey in larger patches to equal that of small patches; thereby controlling for prey density. We found that the intensity at which fish attacked prey (combination of attack rate and duration) was primarily a response to prey density rather than the size of patch they occupied. However, there was evidence that fish spent more time foraging in larger than smaller patches independent of prey density; presumably because of the greater total number of prey available. These experimental observations suggest that fish can distinguish between different notions of prey abundance in ways that enhance their rate of consumption. Although fish may feed in a density dependent manner, a critical issue is whether their rate of consumption outstrips the rate of increase in prey abundance to cause density dependent mortality of prey.     

The effect of prey size and prey density on the functional response,survival, growth and development of a predatory pentatomid bug,Podisus maculiventris say

Two experiments on the nymphal predation of Podisus maculiventris were conducted using Spodoptera litura larvae as prey. First experiment: The predator nymphs divided into three groups were reared individually from second instar to adult in a small vessel. Each nymph in the groups 1, 2 and 3 was allowed to attack the serially growing larvae (these were supplied at the rate of one per day) from 3-, 5- and 7-day old after hatching, respectively. The first prey used for the group 1 was so small that it was not only insufficient to satiate the predator but also was difficult to be searched out. But these disadvantages were soon recuperated due to the rapid growth of the prey and all nymphs could survive to adults. The survival rate of third and fourth instar nymphs in the group 3 was severely affected by vigorous counterattack of older prey larvae. Second experiment: The predator nymphs were individually reared either in a small vessel or in a large one at various rates of food supply (the prey larvae of 7-day old were used). The functional response curves obtained for each instar of the predator took a saturation type within a certain range of the prey density. The saturation level specific to each instar was generally higher for the predator reared in the large vessel than in the small one. The functional response of fourth and fifth instar nymphs was accelerated at a high prey density, viz. 16 larvae per vessel. Even at the low rate of food supply, viz. one larva per day per predator, the predator nymphs could survive to adults, but the size of resultant adults were abnormally small.     

Serially homologous ears perform frequency range fractionation in the praying mantis, Creobroter (Mantodea, Hymenopodidae)

Unlike most praying mantises that have a single region of auditory sensitivity, species in the genus Creobroter have equally sensitive hearing at 2–4 and at 25–50 kHz and and are relatively insensitivity at 10–15 kHz — they have a W-shaped audiogram. Ultrasonic sensitivity originates from an auditory organ in the ventral midline of the metathorax that closely resembles the ear of other mantises. Ablation experiments demonstrate that low frequency sensitivity derives from a serially homologous mesothoracic auditory organ. Extracellular recordings suggest that these two ears operate largely, if not entirely, independently of one another in the thorax. The low frequency response has a longer latency, more action potentials per stimulus, and different patterns of change with increasing SPL than the high frequency response. Separate interneurons mediate responses in the two frequency ranges, but our evidence suggests that they are two serially homologous sets of cells. Neither auditory organ shows any physiological evidence of directional sensitivity. Ultrasound triggers a set of behaviors in flying hymenopodid mantises much like those in other mantises, but the behavioral significance of low frequency hearing in these animals is still unknown.Abbreviations SPL sound pressure level - dB SPL sound pressure level re: 20 Pa - HF high frequency - LF low frequency     

Prey location, recognition and ingestion by the phagotrophic marine dinoflagellate Oxyrrhis marina

The initial ingestion rates of Isochrysis galbana and Dunaliella primolecta by Oxyrrhis populations precultured separately on these phytoplanktonic prey were quantified and related to the chemosensory responses elicited in Oxyrrhis by the filtrate from live and heat killed prey cells. Despite evidence to suggest that Oxyrrhis shows specific distaste towards Isochrysis (but not Dunaliella) such that consumption of N-deplete Isochrysis halted in grazing experiments, positive chemotaxis was observed towards the cell-free filtrate from both species. These results suggest that while tactile cues encountered upon contact with Isochrysis and Dunaliella may enable Oxyrrhis to recognise differences between the two species, the chemosensory responses observed towards dissolved chemical cues derived from potential prey items are non-specific. That chemosensory and ingestion behaviours do not appear to be tightly coupled raises important questions concerning the ecological implications of chemotaxis in Oxyrrhis. Chemotaxis may enhance the overall efficiency of prey detection; however, when confronted with a variety of chemical stimuli (i.e. from a mixed-prey assemblage) Oxyrrhis may be unable to discern the difference between cues that originate from high quality, poor quality (or even toxic) prey items. The positive chemosensory responses observed towards a range of synthetic amino acid, amino sugar and ammonium solutions suggest that chemotaxis could facilitate the detection of solute gradients in prey deplete environments for direct exploitation via osmotrophy. Furthermore, the positive chemotaxis elicited by regenerated ammonium and compounds derived from heat killed conspecifics suggests that Oxyrrhis may release chemical cues which induce cannibalistic behaviour as a ‘life boat mechanism’ when no other suitable (non-self) prey items are available. Further work is required to explore the nature of the chemosensory apparatus and signal transduction pathways that mediate responses to dissolved chemical stimuli in Oxyrrhis and to investigate other sensory mechanisms that enable cells to recognise and differentiate between potential prey items.     

Effect of prey size on attack components of the functional response by Notonecta undulata

The number of encounters per prey, the proportion of encounters resulting in attacks, and the proportion of attacks that were successful were observed while fourth-instar Notonecta undulata nymphs preyed on smaller N. undulata nymphs. While encounters per prey and proportion of encounters resulting in attacks increased with prey size, the proportion of attacks that were successful decreased. The increase in encounter rate per prey was due in part to an increase in the predator's reactive distance to prey as prey size increased. While none of the attack parameters varied significantly with prey density, logarithmic regression of the number of encounters per unit search time on prey density suggested that prey density tends to have a positive effect on encounters per first-instar prey but a negative effect on encounters per second-instar prey. A functional response model is presented that incorporates components of the predator's attack rate as exponential functions of prey density and allows for effects of the time the predator may spend evaluating prey encountered but not attacked and time spent attacking prey not captured. Estimates of the attack parameters derived from the experimental data are used in the model to generate functional response curves for fourth-instar N. undulata preying on first- or second-instar conspecifics. The predicted curve for second-instar prey is typical type II but the curve for firstinstar prey is slightly positively density dependent at low prey densities, i.e., type III.     

Seasonal and taxonomic aspects of the size of arthropods in the tropics and its possible influence on size-selectivity in the prey of a tropical spider community

Summary During one year in Panama the size (=body length) of arthropods (representing the potential prey of a spider community) was determined by sampling with pitfall traps, bowl traps and sweep-netting. Most arthropods found were small and the distribution of their size is considerably skewed to the left (Figs. 1, 2). Pitfall traps and bowl traps yielded similar results but sweep-netting collected larger arthropods and may be size-selective. Although the variation of arthropod size is high, no seasonal influence on body size could be detected in the main prey groups of spiders (Figs. 3, 4). Differences in the size spectra of the actual prey of a tropical spider community are therefore not caused by seasonal variations in the availability of particular prey size classes. Species-specific prey size spectra, however, may be dependent on properties of the webs concerned, microhabitat selection or other, e.g. physiological or ethological differences between the spider species.     

Effects of prey size and foraging mode on the ontogenetic change in feeding niche ofColostethus stepheni (Anura: Dendrobatidae)

The feeding niche ofColostethus stepheni changes during ontogeny. Small individuals eat small arthropods, principally mites and collembolans, and larger frogs eat bigger prey of other types. The shift in prey types is not a passive effect of selection for bigger prey. There is a strong relationship between electivity for prey types and frog size, independent of electivity for prey size. Four indices of general activity during foraging (number of movements, velocity, total area utilized and time spent moving), which are associated with electivity for prey types in adult frogs and lizards, did not predict the ontogenetic change in the diet ofC. stepheni. Apparently, the behavioral changes that cause the ontogenetic change inC. stepheni are more subtle than shifts in general activity during foraging. Studies of niche partitioning in communities of anurans that do not take into consideration ontogenetic changes in diet and seasonal changes in the size structures of populations present a partial and possibly erroneous picture of the potential interactions among species.     

Effects of stream predator richness on the prey community and ecosystem attributes

It is important to understand the role that different predators can have to be able to predict how changes in the predator assemblage may affect the prey community and ecosystem attributes. We tested the effects of different stream predators on macroinvertebrates and ecosystem attributes, in terms of benthic algal biomass and accumulation of detritus, in artificial stream channels. Predator richness was manipulated from zero to three predators, using two fish and one crayfish species, while density was kept equal (n = 6) in all treatments with predators. Predators differed in their foraging strategies (benthic vs. drift feeding fish and omnivorous crayfish) but had overlapping food preferences. We found effects of both predator species richness and identity, but the direction of effects differed depending on the response variable. While there was no effect on macroinvertebrate biomass, diversity of predatory macroinvertebrates decreased with increasing predator species richness, which suggests complementarity between predators for this functional feeding group. Moreover, the accumulation of detritus was affected by both predator species richness and predator identity. Increasing predator species richness decreased detritus accumulation and presence of the benthic fish resulted in the lowest amounts of detritus. Predator identity (the benthic fish), but not predator species richness had a positive effect on benthic algal biomass. Furthermore, the results indicate indirect negative effects between the two ecosystem attributes, with a negative correlation between the amount of detritus and algal biomass. Hence, interactions between different predators directly affected stream community structure, while predator identity had the strongest impact on ecosystem attributes.     

Variation in the effect of profitability on prey size selection by the lacertid lizard Psammodromus algirus

Maximizing the average rate of energy intake (profitability) may not always be the optimal foraging strategy for ectotherms with relatively low energy requirements. To test this hypothesis, we studied the feeding behaviour of captive insectivorous lizards Psammodromus algirus, and we obtained experimental estimates of prey mass, handling time, profitability, and attack distance for several types of prey. Handling time increased linearly with prey mass and differed significantly among prey types when prey size differences were controlled for, and mean profitabilities differed among prey taxa, but profitability was independent of prey size. The attack distance increased with prey length and with the mobility of prey, but it was unrelated to profitability. Thus, lizards did not seem to take account of the rate of energy intake per second as a proximate cue eliciting predatory behavior. This information was combined with pitfall-trap censuses of prey (in late April, mid-June and late July) that allowed us to compare the mass of the prey captured in the environment with that of the arthropods found in the stomachs of sacrificed free-living lizards. In April, when food abundance was low and lizards were reproducing, profitability had a pronounced effect on size selection and lizards selected prey larger than average from all taxa except the least profitable ones. As the active season progressed, and with a higher availability of food, the number of prey per stomach decreased and their mean ize increased. The effect of profitability on size selection decreased (June) and eventually vanished (July–August). This variation is probably related to seasonal changes in the ecology of lizards, e.g. time minimization in the breeding season as a means of saving time for nonforaging activities versus movement minimization by selecting fewer (but larger) prey in the postbreeding season. Thus, the hypothesis that maximizing profitability could be just an optional strategy for a terrestrial ectothermic vertebrate was supported by our data.