Organization of the cuticle of an aquatic fly larva

The aquatic, apneustic larva of the midge, Chironomus riparius, has a very thin (up to 5 micrometers), readily deformable, post-cephalic cuticle. The ultrastructure of this cuticle from newly moulted and older final instar animals, and exuvia shed at pupation, has been examined using routine methods and also after the extraction of proteins with formamide and acetic acid. From the results described, and using established criteria, it is inferred that an exocuticle is present and represents about 25% of the thickness of the mature procuticle, the remainder being endocuticle. Therefore, it would seem that this exceptionally delicate cuticle conforms to the conventional plan of tanned or sclerotized solid cuticles, unlike those reported in the larvae of cyclorrhaphous dipterans such as blowflies or other soft-bodied insects. This is the first account, using experimental techniques, of the fine structure of sectioned cuticle from nematocerous dipteran larvae. It also indicates the value of the exuvium as a source of information about cuticle structure.     

Differential toxicity of leaf litter to dipteran larvae of mosquito developmental sites

The relative toxicity of leaf litter to nematocerous dipteran larvae characteristic of mosquito developmental sites was investigated. Culicidae, Chironomidae, and Simuliidae taxa originating from alpine hydrosystems were tested together with two laboratory nonindigenous culicid taxa. Bioassays indicate that ingestion of 10-month-old decaying leaves from Alnus glutinosa, Populus nigra, and Quercus robur by larvae is more deleterious for Aedes aegypti, A. albopictus, Culex pipiens, Simulium variegatum, and Chironomus annularius than for A. rusticus. Histopathological observations reveal that the midgut epithelium is the main target organ of the toxic effect of dietary leaf litter, which appears to be stronger than that of previously reported tannic acid. There is a general response of the nematocerous larval midgut epithelium to dietary tannins-phenolic compounds: clear cells of the anterior midgut showing symptoms of intoxication before dark cells of the posterior midgut.     

Phytogeny of the nematocerous families of Diptera (insecta)

The relationships of the nematocerous families of Diptera are cladistieally analysed using the parsimony programs PAUP and Hennig86. An extensive review, as well as a data matrix, is presented for 98 almost exclusively morphological characters (larva, 56; pupa, 6; adult, 36). Four infraorders are recognized, viz , Ptychopteromorpha, Culicomorpha, Blephariceromorpha, Bibionomorpha, and a clade containing the 'higher Nematocera' and Brachycera. Traditionally the family Nymphomyiidae or the infraorder Tipulomorpha (=Tipulidae, with or without Trichoceridae) are considered the most basal clade of the extant Diptera. On the basis of our cladistic analysis it is suggested that the Ptychopteromorpha-Culicomorpha clade is the sister-group of all other extant Diptera. We provide evidence that the Axymyiidae are part of a monophyletic Bibionomorpha. The latter infraorder is proposed as the sister-group of the higher Nematocera and Brachycera. We transfer the Tipulidae (Tipulomorpha) to the higher Nematocera, at a position next to Trichoceridae and near the Anisopodidae-Brachycera lineage. Previous hypotheses concerning nematocerous relationships are reviewed.     

Choanoflagellates, choanocytes, and animal multicellularity

Abstract. It is widely accepted that multicellular animals (metazoans) constitute a monophyletic unit, deriving from ancestral choanoflagellate‐like protists that gave rise to simple choanocyte‐bearing metazoans. However, a re‐assessment of molecular and histological evidence on choanoflagellates, sponge choanocytes, and other metazoan cells reveals that the status of choanocytes as a fundamental cell type in metazoan evolution is unrealistic. Rather, choanocytes are specialized cells that develop from non‐collared ciliated cells during sponge embryogenesis. Although choanocytes of adult sponges have no obvious homologue among metazoans, larval cells transdifferentiating into choanocytes at metamorphosis do have such homologues. The evidence reviewed here also indicates that sponge larvae are architecturally closer than adult sponges to the remaining metazoans. This may mean that the basic multicellular organismal architecture from which diploblasts evolved, that is, the putative planktonic archimetazoan, was more similar to a modern poriferan larva lacking choanocytes than to an adult sponge. Alternatively, it may mean that other metazoans evolved from a neotenous larva of ancient sponges. Indeed, the Porifera possess some features of intriguing evolutionary significance: (1) widespread occurrence of internal fertilization and a notable diversity of gastrulation modes, (2) dispersal through architecturally complex lecithotrophic larvae, in which an ephemeral archenteron (in dispherula larvae) and multiciliated and syncytial cells (in trichimella larvae) occur, (3) acquisition of direct development by some groups, and (4) replacement of choanocyte‐based filter‐feeding by carnivory in some sponges. Together, these features strongly suggest that the Porifera may have a longer and more complicated evolutionary history than traditionally assumed, and also that the simple anatomy of modern adult sponges may have resulted from a secondary simplification. This makes the idea of a neotenous evolution less likely than that of a larva‐like choanocyte‐lacking archimetazoan. From this perspective, the view that choanoflagellates may be simplified sponge‐derived metazoans, rather than protists, emerges as a viable alternative hypothesis. This idea neither conflicts with the available evidence nor can be disproved by it, and must be specifically re‐examined by further approaches combining morphological and molecular information. Interestingly, several microbial lin°Cages lacking choanocyte‐like morphology, such as Corallochytrea, Cristidiscoidea, Ministeriida, and Mesomycetozoea, have recently been placed at the boundary between fungi and animals, becoming a promising source of information in addition to the choanoflagellates in the search for the unicellular origin of animal multicellularity.     

黑水虻幼虫及预蛹抗逆性的初步研究

本文初步研究了老熟和预蛹阶段的黑水虻Hermetiaillucens对于酒精毒性、氧胁迫和高渗透压条件下的耐受特性,结果显示黑水虻幼虫和预蛹在三种胁迫因子处于20%～60%(浓度)的较低水平时,其平均死亡率低于20%,表现出良好的抗逆性。高浓度的酒精、矿物油和氯化钠溶液对于黑水虻的生存都会产生负面影响,但是幼虫与预蛹由于其生理形态上的变化,致使其在抗逆性方面表现出显著差异,预蛹阶段的黑水虻对矿物油较为敏感,而幼虫则对高盐环境的耐受性表现较弱。     

Le filet-piège de la larve de Plectrocnemia conspersa (Trichopteres,Polycentropodidae)

Summary The catching-net of P. conspersa larva was studies with particular reference to: structure, building, larval activity inside it and its function in prey catching.The microscopic and macroscopic structure of this catching-net is similar to that Neureclipsis larvae, with an open-ended tunnel and two funnel shaped openings, but its shape is less regular.The building of this net lasts about four to six hours. After a wandering period, the larva first builds the open-ended tunnel and the two openings.Most of the time, the larva lives inside the open-ended tunnel, but it can also crawl into the entrances. Its activity does seem to be related to a diel periodicity.During the catching process, the chief function of the net is detection, after which the prey is taken in the same way as in a spider's web.     

Reciprocal transplantation of wing discs between a wing deficient mutant (fl) and wild type of the silkworm, Bombyx mori

The wingless mutant flügellos ( fl ) of the silkworm lacks all four wings. Although wing discs of the fl seem to develop normally until the fourth larval instar, wing morphogenesis stops after the fourth larval ecdysis, probably caused by aberrant expression of an unidentified factor, referred to as fl . To characterize factor fl , the wing discs dissected from the wild-type (WT) and fl larvae were transplanted into other larvae and developmental changes of the discs were examined. When the wing disc from a WT larva was transplanted into another WT larva and allowed to grow until emergence, a small wing appeared that was covered with scales. Thus, the transplanted wing discs can develop autonomously, form scales and evert from adult skin. The WT wing discs transplanted into the fl larvae also developed at a high rate. However, the fl wing discs transplanted into the WT larvae did not develop during the larval to pupal developmental stages. These data suggest that the fl gene product (factor fl) works in the wing disc cells during wing morphogenesis. Its function cannot be complemented by hemolymph in the WT larva. It is also implied that the level of humoral factors and hormones required for wing morphogenesis are normally maintained in the fl larva.     

The nonfeeding auricularia of Holothuria mexicana (Echinodermata,Holothuroidea)

Among echinoderms, nonfeeding larvae usually are simplified in body shape, have uniform ciliation, and have lost the larval gut. A few species have nonfeeding larvae that express some remnant features of feeding larvae like ciliated bands and larval skeleton or larval arms, but typically their larval mouth never opens and their gut does not function. Still other species have retained the feeding larval form, a functional gut, and can feed, but they do not require food to metamorphose. The present note describes the development of a tropical holothurian, Holothuria mexicana, which hatches as a gastrula that is already generating coelomic structures. A translucent auricularia forms with a mouth that opens but becomes reduced soon thereafter. In form and ciliation this auricularia resembles a feeding larva, but it does not respond to food. A doliolaria forms on day 4 and the pentactula on day 6 post‐fertilization. Further study of this larva and that of its closely related congener, Holothuria floridana, is warranted.     

Antipredator responses to alarm pheromone in groups of young and/or old thrips larvae

Many prey species suffer from different predators in the course of their ontogeny. Hence, the alarm signal a small prey individual sends can have a different meaning than the signal a large prey individual sends, both for small and for large receivers. Larvae of Western Flower Thrips face predators that attack only small larvae, or predators that attack small larvae and large larvae. Furthermore, thrips larvae release a two‐component alarm pheromone, which varies in composition with larval age. Here, we study whether their response to alarm pheromone varies with composition of the pheromone. First, we confirmed that large and small larvae respond when nearby larvae of both sizes were prodded with a brush to induce alarm pheromone excretion. Subsequently, we tested whether thrips larvae of a given size respond differentially to alarm pheromone excreted by a small or large companion larva. We analyzed two types of behavior used in direct defense against a predator and one type of escape response. Only small (not large) larvae attempted to escape more frequently in response to excretions from a large larva. This difference in response could have been due to the alarm pheromone or to the companion larva in the vicinity. We subsequently tested for, but did not find, an effect of size of the companion larva on the behavior of the test larva when exposed to synthetic pheromone mimicking that of a large larva. Finally, we tested how pheromone composition affects antipredator behavior by exposing thrips larvae to synthetic pheromones differing in amount and ratio of the two components. Only for small larvae, we found significant changes in escape behavior with pheromone amount, and a trend with the ratio. Overall, we conclude that small thrips larvae respond differentially to alarm pheromones excreted by small and large larvae and that this differential response is due to differences in pheromone quantity and possibly also quality. Our results suggest that responses to alarm signals can vary with the chemical composition of those alarm signals.     

幼虫密度对斜纹夜蛾抗病能力的影响

为了阐明斜纹夜蛾Spodoptera litura Fabricius幼虫密度对其抗病能力的影响,在室内条件下(温度23℃±1℃,相对湿度75%)对不同幼虫密度(1、2、5、10、15头/皿(直径为12cm))饲养的斜纹夜蛾幼虫抵抗斜纹夜蛾核型多角体病毒侵染的能力及其免疫指标进行了研究。结果表明:幼虫密度对斜纹夜蛾幼虫接种核型多角体病毒后的存活率、存活时间及血淋巴酚氧化酶活性影响显著。随着幼虫密度的增加,接种核型多角体病毒后幼虫的存活率降低,存活时间缩短。当幼虫密度达到15头/皿时,幼虫存活率显著低于其它幼虫密度。不同幼虫密度幼虫的存活时间以1头/皿的最高,15头/皿的最低,且二者之间差异显著。幼虫血淋巴中酚氧化酶活性随幼虫密度的增加而明显降低,当幼虫密度达到5头/皿时,幼虫酚氧化酶活性显著低于1头/皿的。另外,幼虫溶菌酶活性和血细胞总数受幼虫密度影响不显著。不同密度幼虫抗病性的变化与其血淋巴中酚氧化酶活性的变化趋势较为一致。所以斜纹夜蛾幼虫抗病能力的降低可能与幼虫酚氧化酶活性的下降有关。因此,幼虫密度是影响斜纹夜蛾幼虫抗病性变化的重要因子。     

EVOLUTIONARY LOSS OF LARVAL FEEDING: DEVELOPMENT,FORM AND FUNCTION IN A FACULTATIVELY FEEDING LARVA,BRISASTER LATIFRONS

Species with large eggs and nonfeeding larvae have evolved many times from ancestors with smaller eggs and feeding larvae in numerous groups of aquatic invertebrates and amphibians. This change in reproductive allocation and larval form is often accompanied by dramatic changes in development. Little is known of this transformation because the intermediate form (a facultatively feeding larva) is rare. Knowledge of facultatively feeding larvae may help explain the conditions under which nonfeeding larvae evolve. Two hypotheses concerning the evolutionary loss of larval feeding are as follows: (1) large eggs evolve before modifications in larval development, and (2) the intermediate form (facultatively feeding larva) is evolutionarily short-lived. I show that larvae of a heart urchin, Brisaster latifrons, are capable of feeding but do not require food to complete larval development. Food for larvae appears to have little effect on larval growth and development. The development, form, and suspension feeding mechanism of these larvae are similar to those of obligate-feeding larvae of other echinoids. Feeding rates of Brisaster larvae are similar to cooccurring, obligate-feeding echinoid larvae but are low relative to the large size of Brisaster larvae. The comparison shows that in Brisaster large egg size, independence from larval food, and relatively low feeding rate have evolved before the heterochronies and modified developmental mechanisms common in nonfeeding echinoid larvae. If it is general, the result suggests that hypotheses concerning the origin of nonfeeding larval development should be based on ecological factors that affect natural selection for large eggs, rather than on the evolution of heterochronies and developmental novelties in particular clades. I also discuss alternative hypotheses concerning the evolutionary persistence of facultative larval feeding as a reproductive strategy. These hypotheses could be tested against a phylogenetic hypothesis.     

Adult female and neonate larval plant preferences of the generalist herbivore, Epiphyas postvittana

The polyphagous leafroller moth, Epiphyas postvittana, is a pest of many fruit crops in New Zealand. Since the larva is highly mobile, host selection in this insect may involve both the adult female and the larva. In order to test the relative importance of the adult female and the neonate larva in the selection of host plants, the ovipositional preferences of females, and the preferences or acceptances of neonate larvae towards 26 plant species, consisting of 15 plants considered hosts and 11 not considered hosts, were investigated. In the ovipositional tests, the mean preferences of females for hosts and non-hosts were very similar. In contrast, larvae showed a significantly greater mean preference or acceptance towards hosts than to non-hosts, in both choice and no-choice bioassays, respectively. There were highly significant correlations between the preferences and acceptances of larvae for plants in the choice and no-choice tests. In the no-choice tests, there was a highly significant correlation between the acceptances of neonate larvae towards plants after one and three days (i.e., acceptances changed little over time). Moreover, in these no-choice tests, there was a significant negative correlation between larval acceptance at 1 day and larval mortality after 3 days; that is, the less acceptable a given plant at 1 day, the more likely larvae would fail to establish, feed, and survive on it by three days. Female and larval preferences towards the various plants were also negatively correlated. Together, these data suggest that the selection of a plant for the neonate larva to feed on is largely governed by the preferences of the larva, rather than by the preferences of the female. However, selection of a plant for oviposition by the female, may be important in host selection for reasons unrelated to larval preferences, for example, by encouraging dispersal, perhaps to other plant species, of the neonate larvae and thereby decreasing intersibling competition.     

Induction of anhydrobiosis in fat body tissue from an insect

The larva of the African chironomid Polypedilum vanderplanki can withstand complete desiccation. Our previous reports revealed that even when the larva is dehydrated without a brain, it accumulated a great amount of trehalose and successfully went into anhydrobiosis. In this paper we determined the viability after rehydration in tissues from the larvae followed by complete dehydration. Only fat-body tissues that were the main producer of trehalose could be preserved in a dry state at room temperature for an extended period of more than 18 months in a viable form. Thus we have confirmed that the central nervous system is not involved in the induction of anhydrobiosis, even in this complex multicellular organism.     

The biology, egg and larvae of Acaenitus dubitator (Panzer) (Hymenoptera, Ichneumonidae: Acaenitinae)

Acaenitus dubitator (Panzer) is found to be a koinobiont endoparasitoid of the larva of an endophytic beetle, Cleonis piger (Scopoli) (Curculionidae), in Britain, suggesting a similar mode of development for the ichneumonid subfamily Acaenitinae as a whole. The parasitoid can overwinter in its cocoon in one of two ways. Individuals overwintering as essentially unaltered mature larvae do not become adult the following summer, while those that overwinter as morphologically distinct prepupae are committed to pupate and become adult immediately afterwards. The change from mature larva to prepupa takes place in late summer, soon after the time of cocoon formation, but a proportion of mature larvae lie over in the first year, and perhaps subsequently. This appears to be an adaptation to life in a particularly harsh and uncertain environment. The egg, prepupa, and first, second and final instar larvae are described and figured. Previous interpretations of the cephalic sclerites of final instar acaenitines are revised.     

The Larvae of the Mesozoic family Aeschnidiidae and their Phylogenetic Implications (Insecta, Odonata, Anisoptera)

Four giant dragonfly larvae are described from the Lower Cretaceous of China. Owing to the preservation of wing tracheal venation on the larval wing sheaths, they can be identified as the first undoubted larvae of the extinct Mesozoic family Aeschnidiidae. They are ultimate or penultimate male and female specimens, and a younger larva. The female larva has a very long ovipositor sheath. These larvae have an anisopteran anal pyramid and a very particular spoon-shaped labial mask, with a very narrow prementum and large palps with numerous teeth, suggesting possible affinities of the Aeschnidiidae with the Anisoptera Cavilabiata. The positions of other larvae formally attributed to the Aeschnidiidae are discussed, i.e. Nothomacromia sensibilis (Carle and Wighton, 1990), Sona nectes Pritykina, 1986, and the alleged larvae of Hemeroscopus baissicus Pritykina, 1977. They differ greatly from the true Chinese larval Aeschnidiidae, in the labial mask and female ovipositor, even if they show some similarities in the anal pyramid.     

Hatching and earliest larval stages of the priapulid worm Priapulus caudatus

Abstract. Here we describe the hatching and morphology of the earliest larval stages of the priapulid worm Priapulus caudatus for the first time. The hatching larva differs considerably from previously described larvae not only in its general body shape but also in its lack of a proper lorica including the typical lorica tubuli. Furthermore, no mouth opening or pharyngeal teeth have formed as yet, and the number and arrangement of scalids differ from that of later larvae. The hatching larva molts and emerges as the first lorica larva. This larva partially resembles earlier described lorica larvae, but there are a number of important differences; the first lorica larva is smaller, and the mouth opening as well as pharyngeal teeth are still yet to form. The second lorica larva is equipped with four rings of pharyngeal teeth; it shows striking similarity to the previously described larva of P. caudatus , i.e., the larva-type 2 , only differing in the scalid pattern. We conclude that the first two larval stages of P. caudatus have not been described previously. We suggest that discrepancies between the earliest lorica larvae described here and in earlier publications might depend on sub-speciation or ecophenotypic modification of larvae collected from different localities. Our findings highlight the importance of studying the development of non-model organisms such as priapulids under controlled laboratory conditions.     

Eicosanoids mediate Galleria mellonella cellular immune response to viral infection

Nodulation is the predominant insect cellular immune response to bacterial and fungal infections and it can also be induced by some viral infections. Treating seventh instar larvae of greater wax moth Galleria mellonella with Bovine herpes simplex virus-1 (BHSV-1) induced nodulation reactions in a dose-dependent manner. Because eicosanoids mediate nodulation reactions to bacterial and fungal infection, we hypothesized that eicosanoids also mediate nodulation reactions to viral challenge. To test this idea, we injected G. mellonella larvae with indomethacin, a nonsteroidal anti-inflammatory drug immediately prior to intrahemocoelic injection of BHSV-1. Relative to vehicle-treated controls, indomethacin-treated larvae produced significantly reduced numbers of nodules following viral infection (down from approximately 190 nodules/larva to <50 nodules/larva). In addition to injection treatments, increasing dietary indomethacin dosages (from 0.01% to 1%) were associated with decreasing nodulation (by 10-fold) and phenoloxidase activity (by 3-fold) reactions to BHSV-1 injection. We infer from these findings that cyclooxygenase products, prostaglandins, mediate nodulation response to viral infection in G. mellonella.     

Hemps, a novel EGF-like protein, plays a central role in ascidian metamorphosis

All chordates share several characteristic features including a dorsal hollow neural tube, a notochord, a pharynx and an endostyle. Unlike other chordate taxa, ascidians have a biphasic life-history with two distinct body plans. During metamorphosis, the larval nerve cord and notochord degenerate and the pharyngeal gill slits and endostyle form. While ascidians, like other marine invertebrates, metamorphose in response to specific environmental cues, it remains unclear how these cues trigger metamorphosis. We have identified a novel gene (Hemps) which encodes a protein with a putative secretion signal sequence and four epidermal growth factor (EGF)-like repeats which is a key regulator of metamorphosis in the ascidian Herdmania curvata. Expression of Hemps increases markedly when the swimming tadpole larva becomes competent to undergo metamorphosis and then during the first 24 hours of metamorphosis. The Hemps protein is localised to the larval papillae and anterior epidermis of the larva in the region known to be required for metamorphosis. When the larva contacts an inductive cue the protein is released, spreading posteriorly and into the tunic as metamorphosis progresses. Metamorphosis is blocked by incubating larvae in anti-Hemps antibodies prior to the addition of the cue. Addition of recombinant Hemps protein to competent larvae induces metamorphosis in a concentration-dependent manner. A subgroup of genes are specifically induced during this process. These results demonstrate that the Hemps protein is a key regulator of ascidian metamorphosis and is distinct from previously described inducers of this process in terrestrial arthropods and aquatic vertebrates.     

Predatory wasps learn to overcome the shelter defences of their larval prey

Larvae of Epargyreus clarus (Hesperiidae), the silver-spotted skipper, inhabit leaf-and-silk shelters that they construct on their leguminous host plants. In the field, Polistes spp. (Vespidae) wasps land on the shelters, quickly extracting and killing the larvae within. In marked contrast, wasps that emerge from field-collected colonies maintained in the laboratory visit and examine leaflets bearing sheltered caterpillars, but only rarely do they extract and kill the sheltered larvae. To examine whether learning is involved in the development of the ability of Polistes wasps to forage successfully on sheltered E. clarus larvae, we tested the responses of P. fuscatus and P. dominulus wasps to sheltered E. clarus larvae before and after their exposure to unsheltered larvae that were visible either on an opened host-leaf shelter (P. fuscatus and P. dominulus) or on a nonhost leaf in the absence of a shelter (P. fuscatus). After killing and processing an unsheltered larva that was visible on an opened leaf shelter, a majority of foragers subsequently extracted and killed larvae from closed shelters. Wasps that killed and processed an unsheltered larva on a nonhost leaf, on the other hand, generally did not later open shelters. Thus, it seems that experience with an exposed larva in the context of its shelter is necessary for a wasp to be able to prey on sheltered larvae. We conclude that the wasps must learn to associate the taste of the larva with shelter-related cues, such as presence of leaf damage and silk. In nature, this initial exposure may occur when the larva is visible in or near its shelter, perhaps when feeding or constructing a new shelter. Learning opportunities will thus depend on larval density. Our results show that invertebrate predators can learn to overcome their prey's defences, and are therefore able to make use of previously inaccessible prey.     

On interference among larvae of the greasy cutworm,Agrotis ypsilon rott. (lepidoptera: Noctuidae)

Summary The greasy cut worm,Agrotis ypsilon Rott., has a characteristic habit; the larva feeds on the small seedling of the host plant. Studies in a population in outbreak area suggested that the food shortage temporally took place, causing the reduction of weights of pupae and adults. The fact that this species depends on the small seedling as the host seemed to be responsible for this food shortage. However, it was found that the larval crowding also affected the body weight, even if the food was supplied in excess. It was further suggested that the weight reduction is due to the injury caused by the encounter of larvae. The oviposition habit to lay small egg masses and the aggressive behaviour of the larva was considered to be responsible for the uniform distribution at the later larval stage. Probably, the larval dispersion enabled the larva to forage in the most effective way as well as it protected the larva from the attack of another ones. As a result, this would enable the maximum number of individuals to survive. It seemed that the functions noted above did not work in the population being under outbreak condition, because the larval density was unusually high due to mass-invasion of moths.