Are abdominal prolegs serially homologous with the thoracic legs in panorpidae (Insecta: Mecoptera)? embryological evidence

It is a long standing question whether the abdominal prolegs of holometabolous insect larvae are serially homologous with their thoracic legs. The histology and ultrastructure of proleg embryonic development in the scorpionfly Panorpa magna were studied using light and scanning electron microscopy. During the early embryonic development, paired primary abdominal appendages appeared laterally in line with the thoracic legs. Several hours later, a pair of proleg primordia arose along the midventral line on each of the first eight abdominal segments mesial to the primary abdominal appendages, which then ceased to grow and eventually degenerated into flat vestiges. Histological observation showed that the thoracic legs were obviously connected with lateral thoracic muscle cells, whereas the abdominal prolegs resembled secondary outgrowths. No apparent contact was observed between the lumen of abdominal prolegs and the hemocoel. After dorsal closure, each thoracic segment bore a pair of well‐developed five‐jointed legs, whereas the prolegs were unjointed, fleshy structure. The remnants of the primary abdominal appendages could still be clearly seen in the mature embryo. On the basis of the histological and morphological observation of the embryonic development, we confirm that the abdominal prolegs of Panorpidae lack the characters of the primary appendages; hence they are not serially homologous with the thoracic legs. The reasons why the primary abdominal legs are reduced in scorpionflies are briefly discussed. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Comparative embryogenesis of Mecoptera and Lepidoptera with special reference to the abdominal prolegs

The eruciform larvae of holometabolous insects are primarily characterized by bearing a varying number of abdominal prolegs in addition to three pairs of thoracic legs. However, whether the prolegs are evolutionarily homologous among different insect orders is still a disputable issue. We examined the embryonic features and histological structure of the prolegs of the scorpionfly Panorpa byersi Hua and Huang (Mecoptera: Panorpidae) and the Oriental armyworm Mythimna separata (Walker) (Lepidoptera: Noctuidae) to investigate whether the prolegs are homologous between these two holometabolous insect orders. In the scorpionfly, paired lateral process primordia arise on abdominal segments I–VIII (A1–A8) in line with the thoracic legs in early embryonic stages, but degenerate into triangular protuberances in later stages, and paired medial processes appear along the midventral line before dorsal closure and eventually develop into unjointed, cone‐shaped prolegs. Histological observation showed that the lumina of the prolegs are not continuous with the hemocoel, differing distinctly from that of the basic appendicular plan of thoracic legs. These results suggest that the prolegs are likely secondary outgrowths in Mecoptera. In the armyworm, lateral process primordia appear on A1–A10 in alignment with the thoracic legs in the early embryonic stages, although only the rudiments on A3–A6 and A10 develop into segmented prolegs with the lumina continuous with the hemocoel and others degenerate eventually, suggesting that the prolegs are true segmental appendages serially homologous with the thoracic legs in Lepidoptera. Therefore, we conclude that the larval prolegs are likely not evolutionarily homologous between Mecoptera and Lepidoptera. J. Morphol. 277:585–593, 2016. © 2016 Wiley Periodicals, Inc.     

Novel locomotory mechanisms in caterpillars: life-line climbing in Epinotia abbreviana (Tortricidae) and Yponomeuta padella (Yponomeutidae)

Abstract. When alarmed, caterpillars of the tortricid Epinotia abbreviana (Fabricius) and the yponomeutid Yponomeuta padella (Linnaeus) parachute at the end of a single life-line spun out by the head spinnerets.Two methods are employed to climb back up the line.The first makes use of alternate movements of the left and right set of thoracic legs, aided by side-to-side movements of the body.This results in the life-line being wound in around the third pair of thoracic legs.The second method is based on the normal peristaltic rhythm used when walking on a solid substrate.Dorsoventral curling movements gradually transfer life-line from the thorax along the succession of abdominal pro-legs to the claspers at the end of the body.The life-line has similar physical characteristics to spider orb-web silk, and a breaking force equivalent to 5–8 times body weight.The holding force of the claspers when in contact with the ground exceeds the breaking force of the life-line.These studies demonstrate the flexibility of caterpillar locomotion, when confronted with a novel situation.     

The use of honeydew in foraging for aphids by larvae of the ladybird beetle, Coccinella septempunctata L. (Coleoptera: Coccinellidae)

Abstract.  1. To clarify the use of honeydew in foraging for aphids by larvae of the ladybird beetle, Coccinella septempunctata L., searching behaviour of ladybird larvae for Aphis craccivora Koch and Acyrthosiphon pisum Harris and the abundance of honeydew under aphid colonies were examined in laboratory experiments.
2. More larvae responded by climbing the plants with aphids than responded to plants without aphids. When the plants were replaced with sticks, in order to exclude visual and olfactory cues from plants and aphids, more larvae of C. septempunctata climbed sticks above the area that contained honeydew than climbed sticks above the area that did not contain honeydew. Then, ladybird larvae use honeydew as a contact kairomone when foraging for aphids.
3.  Aphis craccivora deposited a larger number of honeydew droplets beneath the plants than did similar numbers of A. pisum. Thus, C. septempunctata larvae licked more frequently the honeydew of A. craccivora than that of A. pisum and spent longer searching on the area containing honeydew of A. craccivora than that of A. pisum . Consequently, a larger number of larvae climbed a stick above honeydew of A. craccivora than that of A. pisum.
4. It may be also considered that C. septempunctata larvae can distinguish honeydew of the two aphid species and respond more strongly to A. craccivora than A. pisum.     

Comparison of lepidopteran larval communities among tree species in a temperate deciduous forest, Japan

Abstract.  1. The enormous diversity of phytophagous insects in forest canopies is hypothesised to be supported by the number of herbivorous species per host tree species or host specificity. It is therefore necessary to examine the effect of host plant species on compositional changes in the herbivore communities.
2. The lepidopteran larval communities were examined in the canopies of 10 tree species in a temperate deciduous forest of Japan. The phylogeny and leaf flush phenology of host plant species were taken into account as factors affecting the herbivore community assembly.
3. Examination of seasonal changes in the larval community structures on each tree species showed that larval species richness, abundance, and evenness decreased significantly from spring to summer. Larval species richness and abundance were characterised by family-level phylogenetic differences among tree species, whereas evenness was determined at a higher taxonomic level.
4. Compositional changes in the larval communities among tree species showed a remarkable pattern, with a phylogenetic effect at a high taxonomic level in spring, similar to evenness, but a phenological effect in summer. This suggests that host specificity could support the lepidopteran larval diversity in spring.
5. These results suggest that the differences in host utilisation of the herbivore, which reflects the phylogenetic effect of the host plants, can be important as a factor affecting the diversity of lepidopteran larval communities in temperate forests.     

Unusual behaviour and morphology of some Rhyacophila Pictet, 1834 caddisfly (Trichoptera: Rhyacophilidae) larvae reflect their ability to use the hyporheic zone

We found that larvae of four Rhyacophila Pictet, 1834 species preferred the hyporheic biotope in rapids and glides, which appears to be an unusual habitat among species in this genus. These species, i.e., Rhyacophila nigrocephala Iwata, 1927, R. nipponica Navas, 1933, R. shikotsuensis Iwata, 1927 and R. kawamurae Tsuda, 1940, belonging to the nigrocephala species group, of which six species occur in Japan. To study the movement behaviour of these species in the hyporheic biotope, we introduced their larvae into a small aquarium with a sandy bottom substrate. The larvae of these species burrowed into the sand bed and moved smoothly through interstices using their forelegs and their highly flexible and elastic abdomens. The larval morphology of these six species differs from that of the representatives of the other species group of Japanese Rhyacophila. They have more slender and flatter head capsules, more elongate abdominal segments, shorter thoracic legs and more slender anal prolegs. These features are adaptations that allow these species to effectively use the hyporheic biotope by enabling them to burrow through the interstices.     

Control of climbing behavior in the cockroach, Blaberus discoidalis. II. Motor activities associated with joint movement

Deathhead cockroaches employ characteristic postural strategies for surmounting barriers. These include rotation of middle legs to re-direct leg extension and drive the animal upward. However, during climbing the excursions of the joints that play major roles in leg extension are not significantly altered from those seen during running movements. To determine if the motor activity associated with these actions is also unchanged, we examined the electromyogram activity produced by the slow trochanteral extensor and slow tibial extensor motor neurons as deathhead cockroaches climbed over obstacles of two different heights. As they climbed, activity in the slow trochanteral extensor produced a lower extension velocity of the coxal-trochanteral joint than the same frequency of slow trochanteral extensor activity produces during horizontal running. Moreover, the pattern of activity within specific leg cycles was altered. During running, the slow trochanteral extensor generates a high-frequency burst prior to foot set-down. This activity declines through the remainder of the stance phase. During climbing, motor neuron frequency no longer decreased after foot set-down, suggesting that reflex adjustments were made. This conclusion was further supported by the observation that front leg amputees generated even stronger slow trochanteral extensor activity in the middle leg during climbing movements.     

Larval morphology of the scorpionfly Dicerapanorpa magna (Chou) (Mecoptera: Panorpidae) and its adaptive significance

Larval morphology can provide valuable characters for taxonomic and phylogenetic analyses of insects and reflect the adaptations to various living habits. Compared with the adult stages, larval study has lagged far behind in Mecoptera. Although several genera of Panorpidae have been studied for their larval stages, the larva of Dicerapanorpa Zhong and Hua, 2013 basically remains unclear. Here the larva of Dicerapanorpa magna (Chou) is described and illustrated in detail for the first time using light microscopy and scanning electron microscopy. The larva is eruciform, with eight pairs of abdominal prolegs in addition to three pairs of thoracic legs, as in other Panorpidae. The most remarkable characteristics of the larvae include a pair of erect subdorsal annulated processes each on abdominal segments I–IX (A1–A9) and a single middorsal annulated process on A10, as well as a pair of prominent compound eyes composed of over 40 ommatidia, which distinguish this genus from other genera of Panorpidae. The annulated processes may have adaptive significance for fossorial and soil-living habits.     

Control of obstacle climbing in the cockroach, Blaberus discoidalis. I. Kinematics

An advantage of legged locomotion is the ability to climb over obstacles. We studied deathhead cockroaches as they climbed over plastic blocks in order to characterize the leg movements associated with climbing. Movements were recorded as animals surmounted 5.5-mm or 11-mm obstacles. The smaller obstacles were scaled with little change in running movements. The higher obstacles required altered gaits, leg positions and body posture. The most frequent sequence used was to first tilt the front of the body upward in a rearing stage, and then elevate the center of mass to the level of the top of the block. A horizontal running posture was re-assumed in a leveling-off stage. The action of the middle legs was redirected by rotations of the leg at the thoracal-coxal and the trochanteral-femoral joints. The subsequent extension movements of the coxal-trochanteral and femoral-tibial joints were within the range seen during horizontal running. The structure of proximal leg joints allows for flexibility in leg use by generating subtle, but effective changes in the direction of leg movement. This architecture, along with the resulting re-direction of movements, provides a range of strategies for both animals and walking machines.     

Visual displays and taxonomic affinities in the parrot genus Trichoglossus

Variation in visual signalling behaviour in ten different species and subspecies was used re-assess taxonomic affinities within the Loriine parrot genus Trichoglossus. Within the genus, visual signals are derived from a range of non-signalling source movements from which a range of both unritualized and ritualized derivative signals have evolved. This evolution of signal movement can be viewed as occurring in a series of stages, beginning with a number of unritualized primary signals and followed by a succession of increasingly ritualized secondary, tertiary and even quaternary derivatives. By treating each primary signal and its subsequent derivatives as a separate, multi-state, taxonomic character, coefficients of similarity were calculated between each of the different taxa. In general, the results of this analysis confirmed taxonomic affinities derived previously from morphological comparisons. In some cases, however, behavioural resemblances suggested plausible alternatives to existing taxonomic theories. The validity of using culturally inherited behaviour patterns for systematic purposes is discussed.     

Additive apportioning of lepidopteran and coleopteran species diversity across spatial and temporal scales in a cool–temperate deciduous forest in Japan

Abstract.  1. Forest entomofauna retain high diversity, and examining beta diversity, or species turnover, among assemblages in a forest community is vital to elucidate the source of this diversity.
2. Under the DIVERSITAS in Western Pacific and Asia–International Biodiversity Observation Year (DIWPA–IBOY) project for simultaneously documenting biodiversity throughout the Western Pacific and Asian Region, 892 lepidopteran species (51 742 specimens) and 355 coleopteran species (11 633 specimens) were collected in 2001 by light traps in a cool–temperate forest in northern Japan.
3. This study evaluated the beta diversity of lepidopteran and coleopteran communities by ecological categories (i.e. trap location, forest strata, sampling days, and months), and assessed the habitat preferences of lepidopteran and coleopteran species.
4.  anova -like additive apportioning models were used to quantify the beta diversity among the categories. The models simultaneously provide assessments of whether species distributions are biased in favour of particular habitats.
5. Significantly high beta diversity was observed among months for both Lepidoptera and Coleoptera. The category of months corresponded fairly well to the number of specialist species detected in the category, although a remarkably large number of significant specialists in Coleoptera were observed on strata.
6. The high beta diversity and number of specialist species among strata in both communities indicate that stratification between canopy and ground, and seasonal variation, played major roles in species composition and the rich entomofauna in the forest. Highly mobile adults were influenced by the vertical spatial scale, as previously suggested for larvae.     

Embryonic development of the scorpionfly Panorpa emarginata Cheng with special reference to external morphology (Mecoptera: Panorpidae)

The Mecoptera are thought to be one of the most primitive groups in the Holometabola, but their embryology is rarely studied. By means of scanning electron microscopy, we studied the external features of the embryo of the scorpionfly Panorpa emarginata in middle and late development. The embryo remains in the superficial position until hatching. Embryonic development can be divided into 10 stages along with the first‐instar larva. The external features are described from the germ band to the first‐instar larva, with special reference to the components and segmentation of the head, the segmentation of abdomen and the formation of abdominal prolegs. Our results confirm that the head consists of an anterior‐most acron and six trunk segments: the labral, antennal, intercalary, mandibular, maxillary, and labial segments. The labrum is confirmed to derive from the paired appendages. Our observations also provide additional direct evidence that the abdominal prolegs are not serially homologous with the thoracic legs. The presence of the eleventh abdominal segment is clarified. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Stable isotope variability of meso-zooplankton along a gradient of dissolved organic carbon

1.  The δ¹³C and δ¹⁵N signatures of zooplankton vary with dissolved organic carbon (DOC), but inconsistent and limited taxonomic resolution of previous studies have masked differences that may exist among orders, genera or species and are attributable to dietary and/or habitat differences. Here we investigate differences among the isotopic signatures of five zooplankton taxa ( Daphnia , Holopedium , large Calanoida, small Calanoida and Cyclopoida) in Precambrian shield lakes with a sixfold range of DOC concentration.
2.  δ¹³C signatures of Daphnia , small calanoids and large calanoids became more depleted with increasing lake DOC, whereas Holopedium and cyclopoid δ¹³C became enriched with increasing DOC concentration.
3.  The variability of δ¹³C and δ¹⁵N isotopic signatures among zooplankton groups was reduced in high-DOC, compared to low-DOC lakes, especially for δ¹³C. Differences in δ¹³C and POM-corrected δ¹⁵N accounted for up to 33.7% and 19.5% of the variance, respectively, among lakes of varying DOC concentration.
4.  The narrow range of signatures found in higher DOC lakes suggests that different taxa have similar food sources and/or habitats. In contrast, the wide range of signatures in low-DOC lakes suggests that different taxa are exploiting different food sources and/or habitats. Together with the variable trends in zooplankton isotopic signatures along our DOC gradient, these results suggest that food web dynamics within the zooplankton community of temperate lakes will change as climate and lake DOC concentrations change.     

Divergent morphologies, performance, and escape behaviour in two tropical rock-using lizards (Reptilia: Scincidae)

The present study quantified microhabitat use, morphology, performance (sprinting, climbing, clinging, and jumping), and escape behaviour of two closely related tropical rock-using lizards. Specifically, the study tested whether: (1) a flatter body and longer limbs enhance performance in rocky habitats; (2) escape behaviour supports predictions based on habitat openness; and (3) there is a trade-off between sprinting and climbing performance. Despite the occupation of generally similar rocky habitats, the habitat of Carlia scirtetis was more open and composed of larger boulders with more regular surfaces, whereas the habitat of Carlia mundivensis was composed of more undergrowth and leaf litter, consisting of smaller boulders with irregular surfaces. The longer legs, flatter body, and greater sprinting and climbing ability of C. scirtetis, supports ecomorphological predictions. By contrast to predictions based on habitat openness, C. scirtetis allowed a potential threat to approach closer and ran further to a refuge than C. mundivensis , suggesting that escape behaviour as determined by performance may be species-specific or decoupled in these two species. The increased sprint speed of C. scirtetis highlighted a performance trade-off, with climbing speed lagging behind that of sprint speed. These results suggest that subtle differences in the structural microhabitat and the degree of habitat openness may ultimately result in substantial differences in morphology, performance, and threat behaviour in closely-related lizard species.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 85–98.     

Comparative flight performance of the seasonal morphs of the cabbage whitefly, Aleyrodes brassicae (Wlk.), in the laboratory

Abstract. 1. The flight capacity of the summer (non-diapausing) and the winter (diapausing) females of A.brassicae was assessed in terms of the climbing flight directed phototactically toward an overhead source of light in a laboratory flight chamber.
2. The summer morph took off only when prodded and its flight bouts were much briefer than those of the winter morph.
3. Immediately on take-off, the summer morph sank in flight and was negatively phototactic in the horizontal plane whereas the winter morph climbed on take-off, being positively phototactic both vertically and horizontally.
4. Three phases of behaviour during flight were observed in the winter morph: a short period of widely fluctuating rate of climb, a long 'cruising' phase of steady climbing, and a final erratic phase when the rate of climb had fallen to and below zero.
5. In general, the mean rate of climb was significantly higher on re-take-off after a landing than immediately before the landing.     

Effects of aggregation pheromone on individual behaviour and food web interactions: a field study on Drosophila

Abstract.  1. The effects of an aggregation pheromone on individual behaviour and food web interactions were investigated in two ecological communities, using Drosophila melanogaster and D. simulans as focal species.
2. Fruit substrates with aggregation pheromone were significantly more attractive to adult D. melanogaster and D. simulans than control fruit substrates, and the response was positively dose dependent. Competing species and natural enemies were also significantly attracted to substrates with the aggregation pheromone of D. melanogaster and D. simulans .
3. Significantly more eggs were deposited on pheromone-treated fruits than on control fruits, and the microdistribution of eggs within fruits was correlated to the microdistribution of the pheromone. The aggregation pheromone induced more females to share the breeding site.
4. The extremely high densities of fruit flies in the large aggregations appeared to reduce the oviposition rate of females. Physical interactions with conspecific and heterospecifics were frequently observed in the aggregations, and often led to patch leaving of the fruit flies.
5. Competition for food among larvae occurred at high densities and parasitism was density dependent. Aggregation pheromones can be directly responsible for these patterns through their effects on the con- and heterospecific behaviour.
6. The combined results show that aggregation pheromones affect a multitude of aspects in the ecology of interacting animals. The importance of incorporating the communication signals in ecological theory of aggregations is discussed.     

The adequacy of collecting techniques for estimating species richness of grassland invertebrates

1.  There is still considerable debate about the most effective methods of sampling invertebrates in monitoring and assessment programmes.
2.  The above-ground invertebrates of a limestone grassland in north-east England were compared between samples from pitfall traps and from a D-vac suction trap combined with a lightweight swish net (SW/DV).
3.  Over 14 000 individuals were captured, with similar numbers in the pitfall and SW/DV samples. A total of 480 species of Hemiptera, Coleoptera, Diptera and Araneae was identified and placed into 14 taxa for further analysis.
4.  The pitfall sample produced species/specimen curves from which it was possible to estimate species richness for all the Coleoptera and Araneae taxa and the calypterate Schizophora. The SW/DV sample was adequate to estimate the species richness of Hemiptera, most Diptera taxa, herbivorous Coleoptera and Linyphiidae.
5.  The proportion of Coleoptera and Araneae taxa that were method-unique was higher in the pitfall sample than the SW/DV sample and vice versa for the Hemiptera and Diptera taxa. Nevertheless, a relatively high proportion of method-unique species of most taxa was found in both sample types, indicating that they can each contribute to assessing species assemblages in grasslands.
6.  Both pitfall traps and SW/DV samples are needed to estimate species richness in grasslands for all taxa except Heteroptera, Homoptera and Lycosidae. Herbivorous Coleoptera and Linyphiidae were collected in numbers adequate for assessing richness in both sample types, but more specimens were required in one or other sample for the remaining taxa.     

Anatomy of external structure and musculature of abdominal segments in the larva of the lepidopteran Pieris rapae crucivora

The external structure of the 1st (AS1) and 4th abdominal segments (AS4) of Pieris rapae is described in terms of pattern of shallow grooves on the cuticle. Both segments have 5 dorsal costae, 3 ventral costae, and an antero-posterior line in addiction to the dorsal and ventral intersegmental folds and a spiracle. AS4 has a pair of prolegs. The musculatures of AS1 and AS4 consist of 44 and 51 muscles, respectively. As in thoracic ones, most attachments of the muscles are located on the cuticular grooves. AS1 and AS4 have similar musculatures. Common to both segments are 89% of AS1 muscles and 84% of AS4 muscles. AS1 has 6 muscles homologous to proleg ones of AS4, including proleg retractors and plantar retractors. Comparison of the musculature of proleg-bearing abdominal segments among different species shows that abdominal musculature of lepidopteran larvae has major homologous and minor specific muscles. From the muscle attachment sites, the role of each muscle is inferred for contraction and bending of the body, lifting up its venter, taking off the crockets from the substrate, and retraction, lateral abduction, and anterior movement of the proleg.     

The size–grain hypothesis: do macroarthropods see a fractal world?

Abstract.  1. In the size–grain hypothesis (a) long legs allow walking organisms to step over gaps and pores in substrate but prohibit them from entering those gaps; (b) the world is more rugose for small organisms; and (c) the relative cost of long legs increases as organisms grow smaller. The hypothesis predicts a positive allometry of leg length ( = mass ^b where b > 0.33 of isometry), a pattern that robustly holds for ants.
2. Toward testing for leg length allometries in other taxa, arthropods were extracted from the Panama leaf litter and measured. Three common taxa (spiders, diplopods, Coleoptera) yielded b s that exceeded 0.33 while three others (Acarina, Pseudoscorpiones, and Collembola) did not. The exponent b tended to increase ( P = 0.06, n = 7) with an arthropod taxon's average body mass.
3. Since leg length in cursorial organisms tends toward isometry in very small and very large taxa (i.e. mammals) this suggests that the size–grain hypothesis may best apply at a transition zone of intermediate body mass: the macroarthropods.
4. Body length was a robust predictor of mass in all groups despite variation in shape.