The larvae of the burrowing mayfly genus Tortopus (Ephemeroptera: Polymitarcyidae)

The larval stage of Tortopus is redescribed based on three species: T. puella from North America, the only species of the genus previously known from larva, and the larvae of T. obscuripennis and T. sarae from South America described here. Generic characters of the larva include: relatively large finger-like gill near base of maxilla, inner margin of mandibular tusks with a subdistal tubercle, straight or weakly convex frontal ridge present between antennae, reduced unilamellated gill on abdominal segment I. Additionally the male imagines of both Neotropical species are described for the first time, and T. obscuripennis is recorded from Bolivia. Diagnoses, SEM photographs, and illustrations are given for the new stages described and for the identification of the three Tortopus species known as larvae.     

Performance and impact of the biological control agent Xubida infusella (Lepidoptera; Pyralidae) on the target weed Eichhornia crassipes (waterhyacinth) and on a non-target plant, Pontederia cordata (pickerelweed) in two nutrient regimes

Xubida infusella (Walker) (Lepidoptera: Pyralidae) is potentially a useful biological control agent targeting Eichhornia crassipes (waterhyacinth) in the USA but many regions infested with waterhyacinth are also inhabited by an alternative native host, Pontederia cordata (pickerelweed). Experiments were conducted in Australia to assess the impact of X. infusella on pickerelweed compared to waterhyacinth where both these plants were available and X. infusella had already been released. Overall X. infusella had a greater impact on pickerelweed than on waterhyacinth. More than one larva per plant was required to reduce the total shoot dry weight of waterhyacinth but only one larva per plant reduced the total shoot dry weight of pickerelweed. Insect feeding caused the number of secondary shoots (daughter plants) of pickerelweed to double whereas the number of daughter plants produced by waterhyacinth remained unchanged. We suggest this indicates a considerable impact on pickerelweed rather than effective compensation for insect damage because the shoots produced were very small. Waterhyacinth produced a constant number of daughter plants when fed on by up to three larvae per plant. Higher nitrogen status of both species of host plant increased the rate of larval development and pupal weight of X. infusella. The weight and fecundity of X. infusella reared on pickerelweed were lower than those reared on waterhyacinth but large numbers of progeny were produced on both plant species. This experiment demonstrates a considerable impact of X. infusella on pickerelweed suggesting this plant is at risk from this agent if released in the USA where pickerelweed is present. The considerable impact on waterhyacinth demonstrates the potential for this insect to contribute to waterhyacinth control in countries where risk assessment favours release.     

Larvae of the deep-sea Nematocarcinidae (Crustacea: Decapoda: Caridea) from the Southern Ocean

The early larval stages of the deep-sea Nematocarcinidae, Nematocarcinus longirostris Bate, 1888, from the south-western Atlantic Ocean, and N. lanceopes Bate, 1888, from the high Antarctic Weddell Sea, were obtained from plankton catches, described and illustrated. Furthermore, field collected larvae of N. lanceopes were compared with larvae hatched and reared under constant laboratory conditions. The morphology of larvae in both species clearly indicates a planktotrophic and extended mode of larval development. This is an outstanding feature in deep-sea and especially in high-latitudinal caridean shrimp species, and the consequence of such reproductive trait for life history adaptations to both deep-sea and polar environments is discussed.This revised version was published online in November 2004 with corrections to the figure part lettering.     

Endoparasitoids of larval Anomis privata (Lepidoptera: Noctuidae), major pest of Hibiscus syriacus (Columniferae: Malvaceae)

Endoparasitoids of Anomis privata larvae include five species in three families of two orders. In this work, two species of Hymenoptera Braconidae (Cotesia sp., Microplitis sp.), one species of Ichneumonidae (Mesochorus vittator) and two species of Diptera Tachinidae (Exorista (Podotachina) sorbillans, Timavia amoena) were investigated. Of the 261 larvae of A. privata examined, 32 had a parasite, so the rate of parasitism was 12.26%. Parasitism by taxon was the highest, at 10.35% (27 individuals), in Cotesia sp. in Hymenoptera Braconidae. Parasitoids of Braconidae and Ichneumonidae were larval parasitoids. A parasitic insect of Tachinidae was a larva–pupal parasitoid. Solitary parasitoids included Microplitis sp. in Braconidae and E. sorbillans in Tachinidae. Gregarious parasitoids included Cotesia sp. in Braconidae, M. vittator in Ichneumonidae and T. amoena in Tachinidae. There was also a multiparasitoid (T. amoena) and two superparasitoids (Cotesia sp., M. vittator). A larva of A. privata sought feed even after it was parasitized every parasitoid investigated in this study, so five species of parasitoids were all koinobiont.     

Studies on the characteristics of parasitism ofPlutella xylostella (Lep.: Plutellidae) by a larval parasiteDiadegma semiclausum (Hym.: Ichneumonidae)

Laboratory experiments were conducted to study the characteristics of parasitism of diamondback moth,Plutella xylostella (L.), a worldwide pest of crucifers, by a larval parasite,Diadegma semiclausum Hellén. Results showed that the greater the host larval density, the lesser was the percentage of larvae parasitized. The area of discovery decreased with increasing parasite density, but k-value remained practically unchanged. Parasitism reduced food consumption in parasitised larvae. No difference was observed in duration of larval instars between the parasitized and healthy larvae. As the frequency of superparasitism increased, the proportion of production of female progeny also increased. Presence of parasite larva within the host larva deterred superparasitism greater than the presence of parasite egg. Presence of larva of eitherD. semiclausum or another larval parasite,Cotesia plutellae Kurdjumov, within host larva deterred multiparasitism by either species greater than the presence of egg of either parasite species.Diadegma semiclausum was much more aggressive thanC. plutellae in parasitisingP. xylostella larvae.     

Larval host plants of two lizard beetles in the genus Tetraphala (Coleoptera,Erotylidae, Languriinae,Languriini) from Taiwan,with a host plant list of Languriini

Lizard beetles (Erotylidae, Languriinae, Languriini) are known as stem borers of plants and contain agricultural pests and endangered species, but their species–host plant associations have been poorly documented. Here we investigated the larval host plants of two species of the genus Tetraphala Strum, T. collaris (Crotch) and Tetraphala sp. occurring in Taiwan. Females of T. collaris excavated living leafstalks and stems of the herbaceous dicot, Sambucus chinensis (Adoxaceae) using their mandibles for oviposition. We observed the eggs and early-instar larvae inside and nearby oviposition holes and late-instar larvae inside stems, suggesting that T. collaris uses living leafstalks and stems of S. chinensis as oviposition substrate and the larvae tunnel into stems with feeding on the tissues. Similarly, females of Tetraphala sp. excavated living leafstalks of the fern, Pteris wallichiana (Pteridaceae) using their mandibles for oviposition. We observed the eggs and early-instar larvae inside and nearby oviposition holes. When reared in laboratory, a larva reached adulthood inside the leafstalk. These results indicated that Tetraphala sp. uses living leafstalks of Pt. wallichiana as oviposition substrate and the larvae complete their development within. This study revealed that the genus Tetraphala contains both fern- and dicot-users during larval period. Further study is needed to clarify the evolutionary process of host plant use of languriines. Additionally, the host plant list of Languriini is provided.     

Inferring larval taxonomy and morphology in Maladera species (Coleoptera: Scarabaeidae: Sericini) using DNA taxonomy tools

Based on a comparative molecular study of scarab chafers we matched adult and larval instars to identify and describe unknown larvae of Sericini. Here, we use for the first time a two‐fold DNA taxonomy approach based on: (i) mitochondrial and nuclear DNA markers of a local sample (from Nepal) of adults and larvae, in combination with character and tree‐based species delimitation methods; and (ii) a global search of cytochrome c oxidase subunit I (cox1) sequences with GenBank data. In the latter analysis we used a sequence of a specimen that resulted in the first analysis conspecific with the larvae of Maladera affinis (Blanchard) as the query sequence in GenBank, and checked in a minimum evolution tree whether larva–adult matches from the local approach were altered through interference with other taxa of the worldwide database. Both approaches unambiguously identified the unknown larvae as belonging to M. affinis and Maladera cardoni (Brenske). Based on this robust framework of taxonomic identification we could associate names to the larval morphology of the third larval instar of these two Nepalese Maladera species, which are both known for their economical importance in agriculture. They are described here in detail and are compared with known related taxa, especially with Maladera castanea (Arrow).     

Contrary food requirements of the larvae of two Curtonotus (Coleoptera: Carabidae: Amara) species

The larvae of carabids that are granivorous as adults can be granivorous, omnivorous or carnivorous. The differences in larval food preferences of Amara aulica and Amara convexiuscula, two closely related species of the subgenus Curtonotus, were studied. Survival and duration of development of non‐diapausing first and second instar larvae were compared. The third instar larvae were not studied because they go into diapause prior to pupation. The larvae were fed diets consisting of insect larvae (Tenebrio molitor), seeds of Artemisia vulgaris, Tripleurospermum inodorum, Urtica dioica or Cirsium arvense and a mixed diet consisting of insect larvae and seed. Larvae of A. aulica required seeds for successful development, whereas those of A. convexiuscula developed on both an insect diet and seeds of Artemisia. The difference in larval food requirements facilitates the coexistence of these closely related species, which frequently share the same habitat.     

Adenanthera pavonina trypsin inhibitor retard growth of Anagasta kuehniella (Lepidoptera: Pyralidae)

Anagasta kuehniella is a polyphagous pest that feeds on a wide variety of stored products. The possible roles suggested for seed proteinase inhibitors include the function as a part of the plant defensive system against pest via inhibition of their proteolytic enzymes. In this study, a trypsin inhibitor (ApTI) was purified from Adenanthera pavonina seed and was tested for insect growth regulatory effect. The chronic ingestion of ApTI did result in a significant reduction in larval survival and weight. Larval and pupal developmental time of larvae fed on ApTI diet at 1% was significantly longer; the larval period was extended by 5 days and pupal period was 10 days longer, therefore delaying by up to 20 days and resulting in a prolonged period of development from larva to adult. As a result, the ApTI diet emergence rate was only 28% while the emergence rate of control larvae was 80%. The percentage of surviving adults (%S) decreased to 62%. The fourth instar larvae reared on a diet containing 1% ApTI showed a decrease in tryptic activity of gut and that no novel proteolytic form resistant to ApTI was induced. In addition, the tryptic activity in ApTI ‐fed larvae was sensitive to ApTI. These results suggest that ApTI have a potential antimetabolic effect when ingested by A. kuehniella. © 2010 Wiley Periodicals, Inc.     

Ecological and morphological characteristics of endoparasitoids on Elcysma westwoodii (Vollenhoven) (Lepidoptera: Zygaenidae)

Six species of insect endoparasitoids were identified from Elcysma westwoodii, which is the most damaging lepidopteran pest of Prunus yedoensis. From Hymenoptera, two species were identified: a species in Braconidae and Charops striatus in Ichneumonidae. From Diptera, there were four species in Tachinidae: Compsilura concinnata, Exorista sp., Pales sp. and Tachinidae spp. The parasitic ratio was 4.86% (45 of 926 larvae). The hymenopterans were parasitic on 31 individuals of E. westwoodii (68.9%) and the dipterans were parasitic on 14 individuals (31.1%). It was found that parasitoids from the larvae of E. westwoodii were all either endoparasitoids or larval parasitoids. However, Exorista sp. of Tachinidae was found to be either a larval parasitoid or larval-pupal parasitoid. Additionally, all the identified parasitoids were solitary parasitoids, as only one parasite occurred in a larva of E. westwoodii. Because the larva of E. westwoodii eats and molts after it is parasitized, all the parasitoids were identified as koinobionts. There were no big differences in morphological characteristics and life histories between C. striatus and C. concinnata. However, for Exorista sp. and Pales sp., males took 3–5 days longer to emerge from their pupae and had remarkably longer body lengths than females.     

Adaptations to captive breeding of the longhorn beetle Morimus funereus (Coleoptera: Cerambycidae); application on amylase study

Captive breeding has been suggested as a method of conservation for many vertebrates, and is increasingly being proposed as a strategy for invertebrates. In this study, the growth, development and fertility of adults of the vulnerable cerambycid Morimus funereus reared in captivity are examined. Two oviposition cycles; from May to September and from January to March were studied and larvae from wild adults and from the progeny of captive adults (second generation larvae) were examined. Five to 12 instars were observed during larval development. Larval development was completed in 218 days (average) for the progeny of wild adults with an average mortality rate of 10.3% and in 226 days (average) for larvae from captive adults with mortality rate of 34.9%. First generation larval body weights were disparate during development, while second generation larvae had similar weights with no significant differences. In this study we have tested the potential of captive breaded M. funereus larvae as a model for investigation of digestive enzymes. Amylase from the midgut of larvae reared under laboratory conditions showed twofold higher specific activities with a decreased number of isoforms expressed, as compared to the enzyme from field-collected larvae. Captive breeding of M. funereus can be used in the future as a part of an effective conservation strategy for this rare insect species.     

Association of insect life stages using DNA sequences: the larvae of Philodytes umbrinus (Motschulsky) (Coleoptera: Dytiscidae)

Abstract. Insect life stages are known imperfectly in many cases, and classifications are based often on only one or a few semaphoronts of a species. This is unfortunate as information in alternative life stages often is useful for scientific study. Although recent examples of DNA in taxonomy have emphasized the identification of indistinguishable species, such sequence data facilitate the association of life history stages and hold considerable promise in phylogenetic analysis, evolutionary studies, diagnostics, etc. These concepts are discussed here and an example is provided from diving beetles (Dytiscidae: Coleoptera). Three unknown larval specimens of an apparent species of Laccophilinae collected in Namibia were associated with the species Philodytes umbrinus (Motschulsky) using DNA sequence data. An 806-bp portion of the gene cytochrome oxidase I was sequenced from the unknown larvae. Several identified adult specimens of species of Laccophilinae from Namibia were also sequenced, including two P. umbrinus specimens and specimens from four Laccophilus Leach species. Additional species of Laccophilus from other areas of the world also were sequenced, as were specimens of Agabetes acuductus (Harris), Australphilus saltus Watts, Neptosternus boukali Hendrich & Balke and a species of Laccodytes Régimbart. Parsimony analysis resulted in two most parsimonious trees with the unknown larva unambiguously resolved in a group with both adult specimens of P. umbrinus (bootstrap value = 100%). The average pairwise p-distance between the unknown larva and adult P. umbrinus specimens averaged 0.09% (0–0.14%), compared with an average divergence between other conspecifics in the analysis of 0.24% (0–0.82%) and an overall average divergence between species of 13.49% (1.90–19.86%). Based on this, the unknown larvae were assigned to P. umbrinus. The larvae are diagnosed and described and their relationship with other Laccophilinae is discussed.     

The response of Hyssopus pallidus to hosts previously parasitised by Ascogaster quadridentata: heterospecific discrimination and host quality

1. Two basic tenets of competition among parasitoids, that taxonomically distinct parasitoids are unable to discriminate against hosts that have previously been attacked by a competitor and that previous parasitism reduces the quality of a host, were tested by monitoring the oviposition response of Hyssopus pallidus, a gregarious ectoparasitoid, to healthy codling moth larvae and codling moth larvae that had previously been parasitised by a solitary endoparasitoid, Ascogaster quadridentata. 2. Hyssopus pallidus accepted both categories of host larva for oviposition when its competitor was constrained as a first‐instar larva by the diapause development of its host, but discriminated against previously parasitised host larvae when its competitor was present as a larger larva in a non‐diapausing host. 3. Hyssopus pallidus distinguished between the two categories of host larva by allocating twice as many eggs to host larvae previously parasitised by A. quadridentata, a response that was not influenced by previous oviposition experience. 4. The larger clutch sizes allocated to previously parasitised host larvae produced twice as many female progeny, each of a typical size, such that the total biomass was twice that produced from the smaller clutches laid on healthy host larvae. Possible confounding influences of host age and diapause are discounted. 5. These results demonstrate that interspecific discrimination does occur in H. pallidus and that host quality can be improved through previous parasitism by an endoparasitoid. Although interspecific discrimination appears rare among insect parasitoids, it may have been overlooked among ectoparasitoids and requires examination of the fitness consequences of interspecific interactions to clarify its adaptive significance.     

Plant abandonment behavior and fitness of monarch larvae (Danaus plexippus) is not influenced by an intraspecific competitor

Integrating aspects of larval lepidopteran behavior that enhance survival into conservation plans could increase the overall impact of the efforts. We previously recommended that where possible, maintaining 2–4 ramets of closely-spaced common milkweed would support the development of at least one monarch through pupation, based on a seemingly innate behavior in which monarch larvae (Danaus plexippus) abandon their natal milkweed ramet (Asclepias sp.). Here, we explored the impact of intraspecific competition on larval ramet abandonment behavior and fitness of monarch larvae in small artificial milkweed patches.

We observed larvae reared under direct and indirect intraspecific competition, and larvae reared alone.

We found no influence of intraspecific competition; however, our study provides further support that milkweed ramet abandonment is a seemingly innate behavior. This behavior occurs before all of the available leaf biomass on a ramet is consumed and prior to the pre-pupal wandering stage.

Implications for insect conservation: Results from our study suggest that in the absence of predation, parasitism, and interspecific competition, and when sufficient plant biomass is present to support larval development, the presence of an intraspecific competitor does not influence larval behavior or fitness. Based on milkweed ramet abandonment behavior, we continue to suggest maintaining small patches of 2–4 milkweed ramets when possible.

     

Oviposition,larval survival and leaf damage by the willow leaf blotch miner,Micrurapteryx salicifoliella,in relation to leaf trichomes across 10 Salix species

1. Foliar trichomes clearly reduce chewing damage and efficiency of movement by some insect herbivores, but the effect of trichomes on insect oviposition is less well characterised. Trichomes are likely to have particularly strong, negative effects on species that require secure attachment of the egg to the leaf epidermis for successful transition to the feeding stage – a group that includes many leaf mining insects. 2. One such species, Micrurapteryx salicifoliella, must initially enter leaf cells directly from an egg adhered to the cuticle, but later instars can move between leaves and initiate new mines from the leaf exterior. 3. Natural patterns of occurrence by M. salicifoliella were quantified on 10 sympatric Salix species varying in trichome expression to test whether trichomes were associated with reduced oviposition, larval survival and leaf damage. 4. Mean egg density and leaf mining damage were negatively related to mean trichome density across Salix species. Survival of M. salicifoliella from egg to pupa was positively related to trichome density, suggesting that initiation of new mines by late‐instar larvae was not adversely affected by trichomes. There was no evidence that trichomes benefited leaf miner larvae indirectly by decreasing density‐dependent mortality; rather, the positive relationship between trichome density and larval survival may reflect less effective chemical defence by Salix species expressing high trichome density. 5. The results suggest that foliar trichomes serve as an effective defence against M. salicifoliella by deterring oviposition, but do not reduce the survivorship of those individuals that successfully transition from egg to larva.     

HYBRIDIZATION IN LEOPARD FROGS (RANA PIPIENS COMPLEX): LARVAL FITNESS COMPONENTS IN SINGLE-GENOTYPE POPULATIONS AND MIXTURES

Recognizing the predominant mode of selection in hybrid systems is important in predicting the evolutionary fate of recombinant genotypes. Natural selection is endogenous if hybrid genotypes are at a disadvantage relative to parental species independent of environment. Alternatively, relative fitness can vary in response to environmental variation (exogenous selection), and hybrid genotypes can possess fitness values equal to or greater than that of parental species. I investigated the nature of natural selection in a leopard frog hybrid system by rearing larvae of hybrid and parental genotypes between Rana blairi and R. sphenocephala in 1000-L outdoor experimental ponds. Three hybrid (F₁, backcrossj [B₁], backcross₂ [B₂]) and two parental (R. blairi [BB] and R. sphenocephala [SS]) larval genotypes were produced by artificial fertilzations using adult frogs from a natural population in central Missouri. Resultant larvae were reared in single-genotype populations and two-way mixtures at equal total numbers from hatching to metamorphosis. In single-genotype ponds, F₁ hybrid larvae had highest survival and BB were largest at metamorphosis. When F₁ and SS larvae were mixed together, F₁ hybrids had reduced survival and both F₁ and SS larvae metamorphosed at larger body masses than when reared separately. When mixed, both B₁ and SS larvae had shorter larval period lengths than when reared alone. Higher proportion of B₁ metamorphs were produced when larvae were mixed with either parental species than when reared alone. Larval fitness components as measured by survival, body mass at metamorphosis, proportion of survivors metamorphosing, and larval period length for B₂ hybrid and BB larvae were similar in single-genotype populations and mixtures. Comparison of composite fitness component estimates indicated hybrid genotypes possess equivalent or higher larval fitness relative to both parental species for the life-history fitness components measured. Despite reduced survival of F₁ hybrids in mixtures, backcross-generation hybrid genotypes demonstrated high levels of larval growth, survival, and metamorphosis in mixtures with parental species. Consequently, this study suggests natural hybridization and subsequent backcrossing between R. blairi and R. sphenocephala can produce novel and relatively fit hybrid genotypes capable of successful existence with parental species larvae. Thus, the evolutionary fate of hybrid and parental genotypes in this system may be influenced by exogenous selection mediated by genotypic composition of larval assemblages.     

Density‐dependent polyphenism and baculovirus resistance in teak defoliator,Hyblaea puera (Cramer)

1. Field populations of the teak defoliator larvae, Hyblaea puera Cramer exhibit colour polyphenism under different population densities: greyish‐green with black‐ and orange‐coloured dorsal bands in low‐density endemic populations and uniformly black or intermediate colour during high‐density population. 2. The density dependence of colour polyphenism was confirmed by field monitoring of H. puera populations during 2008–2010. 3. The above findings were later substantiated by rearing H. puera larvae under different densities (i.e. solitary and crowded in the laboratory). Ninety one per cent of the solitary reared laboratory population developed bright coloration whereas, 92% of the group reared larvae turned to black. Eight per cent of larvae from both the rearing densities were of intermediate colour. 4. Density‐dependent resistance build‐up against H. puera nucleopolyhedrovirus by H. puera were tested using the fifth instar larvae. The results showed three‐fold increase of median lethal dose (LD₅₀) value for the group reared larvae (5332 polyhedral occlusion bodies/larvae) compared to the solitary reared ones (1727 polyhedral occlusion bodies/larva) and also significant difference for the mean time to death (3.6 and 3.3 days, respectively). 5. The study revealed the strong influence of larval density on H. puera larval melanism and resistance build‐up against H. puera nucleopolyhedrovirus.