Why the larch bud-moth cycle collapsed in the subalpine larch-cembran pine forests in the year 1990 for the first time since 1850

The larch bud-moth cycle has been observed in the sub-alpine larch-cembran pine forests 16 times since 1850. Infestation is easily recognized by the characteristic red-brown discoloration of the larch crowns due to the wasteful feeding of the bud moth larvae. The heaviest defoliation recurs at intervals of 8.47±0.27 (SE) years, and the larval density per kilogram of larch branches varies more than 10000-fold over four or five generations. The basic regulatory mechanism for this cycle is the induced change in food quality for the two or more subsequent larval generations. Defoliation functions as a negative feedback mechanism acting on larval density. In 1989 local discoloration in the Upper Engadine valley was observed in the usual first focus. In 1990 and 1991, however, instead of the expected widespread defoliation damage, larval densities decreased drastically. Based on extensive field data collected from 1961 to 1991 on the development and the survival of the bud moth (Zeiraphera diniana Gn.) and the phenology of the host, Larix decidua L., this paper shows the effect of weather on survival in the egg stage and on the coincidence of larval hatching with the sprouting of the larch. It is shown that the winter and spring weather conditions in 1989–1991 were conducive to unusually high egg mortality. Since these conditions occurred in three successive generations, population growth was effectively reduced and the cycle collapsed prematurely. Thus the rather persistent cyclicity of the larch-larch bud-moth system was disturbed by weather conditions with a very low probability of occurrence, but due to the inherent high resilience of the system, the next population peak with visible defoliation is expected to occur 1996/1997, provided that the weather conditions return to the climatic standard.     

Linking ‘10‐year’ herbivore cycles to the lunisolar oscillation: the cosmic ray hypothesis

The mean population cycle periods of the snowshoe hare Lepus americanus, the larch budmoth Zeiraphera diniana, and the autumnal moth Epirrita autumnata are 9.3 years, similar to the lunar nodal phase cycle. When the full/new Moon is situated close to the ecliptic plane at solstice, it interacts more strongly with the magnetosphere, which plays a crucial role in protecting the Earth against ionizing cosmic ray particles. Ionization by cosmic rays induces protein mobilization in plants, which may increase forage quality for herbivores. Series of hare/lynx population indices from Canadian provinces and an autumnal moth series from Fennoscandia correlated with the lunar nodal phase cycle with different time lags. Both the time lag and the impact of an active sun, which increases solar energetic particles, but decreases galactic cosmic ray particles, were related to radiation or the distance from the auroral oval. These insights improve our ability to explain and understand population peaks, and should also motivate further studies on the effects of cosmic rays on plant chemistry and herbivore performance.     

Effets deBacillus thuringiensis chez une population de Tordeuse grise du mélèze,Zeiraphera diniana [Lep.: Tortricidae] en culmination gradologique

Résumé En 1971, fut dispersée par hélicoptère sur 670 ha de mélèzin de la commune de Montgenèvre (Hautes-Alpes), une suspension de spores et de cristaux toxine deBacillus thuringiensis Berliner, sérotype 1, chez une population deZeiraphera diniana Guénée ayant atteint la culmination gradologique. L'application qui fut effectuée au 3^e stade larvaire du ravageur avant l'apparition des dégats eut une efficacité immédiate moyenne de 80–83%. Cette réduction massive de la population larvaire eut pour conséquence de contenir les effectifs de la Tordeuse du mélèze en dessous de la limite supérieure du niveau de population supportable, pendant la phase dynamique de la gradation. Les dégats ont été pratiquement supprimés sur l'ensemble de la forêt traitée pour la durée de la gradation. Aucune nuisance consécutive à l'intervention bactérienne n'a été constatée.

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Summary In 1971, at Montgenèvre (Hautes-Alpes, France), a suspension of spores and cristals ofBacillus thuringiensis;Berliner Serotype 1, was applied with an helicopter on 670 ha of larch to control a population ofZeiraphera diniana Gn. in outbreak. The application was made on the 3rd instar, before the insect had cause any damage, and showed 80–83% of immediate efficiency. The large scale reduction of the larval population resulted in keepingZ. diniana population below the economic injury level during the dynamic period of gradation. There was practically no damage in the treated forest during this time. No harmful effect due to the bacterial intervention has been observed.

     

Influences of host size and host species on theinfectivity and development of Heterorhabditismegidis (strain NLH-E87.3)

The infectivity, time to first emergence of infective juveniles (IJs), total number of IJs per insect and IJs body length of the entomopathogenic nematode Heterorhabditis megidis (strain NLH-E87.3) after development in larvae of two insect hosts, Galleria mellonella (greater wax moth) and Otiorhynchus sulcatus (vine weevil) was studied. At a dose of 30 IJs, larvae of G. mellonella show to be significantly more susceptible than O. sulcatus larvae. At a dose of one IJ, vine weevil larvae were more susceptible. The number of invading infective juveniles (IJs) increased with host size while the host mortality at a dose of one IJ decreased with the increase of host size. Time to first emergence was longer at a dose of one IJ per larva and increased with the increase of host size in both insect species. Reproduction of IJs differed between host species, host sizes and doses of nematodes. Generally, the IJs body size increased with an increasing host size. The longest infective juveniles were produced at the lowest IJ doses. Results are discussed in relation to the influence of different host species and their different sizes on the performance of H. megidis (strain NLH-E87.3) as a biological control agent.     

HOST-INDUCED ASSORTATIVE MATING IN HOST RACES OF THE LARCH BUDMOTH

Abstract The likelihood of sympatric speciation is enhanced when assortative mating is a by‐product of adaptation to different habitats. Pleiotropy of this kind is recognized as important in parasites that use their hosts as a long‐range cue for finding mates, but is generally assumed to have limited applicability for most other organisms. In the larch budmoth, Zeiraphera diniana (Lepidoptera: Tortricidae), sympatric host races feed on larch or pine. Zeiraphera diniana females attract males (call) by releasing host‐independent long‐range pheromones. Pheromone composition differs strongly between host races, but we show in an experimental field study that cross‐attraction can occur at a rate of 0.03–0.38. Cross‐attraction to larch females increases when they call from neighborhoods (8‐m radius) rich in pine or from pine trees. Cross‐attraction to pine females similarly increases when calling from neighborhoods rich in larch, but there is no significant effect of calling substrate. Males, as well as females, of this species preferentially alight on their own host, and in neighborhoods where their own host is common. This effect of tree species and host neighborhood on assortative mating is therefore due, at least in part, to the numbers of males of each host race present within approximately 200 m² surrounding the female. This proximity effect is enhanced by the clumped distributions of the hosts themselves. Host chemistry might also affect pheromone production and/or response directly, but we have evidence neither for nor against this. This work provides empirical evidence that host adaptation has a pleiotropic effect on assortative mating in a species with host‐independent long‐range mating signals. Sympatric speciation via pleiotropy between ecological traits and assortative mating may thus be more common than generally supposed: Clumped resource distributions and habitat choice by adults are widespread.     

Efficacité des parasites Eulophides deZeiraphera diniana Guenée [Lep., Tortricidae] en fonction de l'état de l'hote

Résumé Un système de notation se référant à une ?gradation-type? (fig. 1) est proposé pour les espèces à gradations périodiques ou occasionnelles. La vitalité de plusieurs souches deZ. diniana a été estimée dans un même biotope et sous des conditions identiques pour diverses densités larvaires. Le critère utilisé pour mesurer cette vitalité était le nombre moyen de pousses de mélèze attaquées par chenille. Dans cet essai, la vitalité des chenilles a varié avec leur stade gradologique (fig. 1), la densité larvaire influenćcant aussi cette relation (tabl. 3). Dans les conditions des deux séries d'essais dont on dispose actuellement, le parasitisme causé par les Eulophides augmente avec la densité des h?tes jusqu'à un certain niveau (tabl. 4). On suppose que la faible capacité de recherche des pondeuses limite le développement de l'importance des Eulophides par basse densité, d'autres facteurs intrinsèques limitant leur efficacité lors de la culmination.

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Summary A method of notation based on a “typical gradation” hereby defined (fig. 1) is proposed for species with periodical or irregular population dynamics. The vitality ofZ. diniana from different areas could be estimated in one biotop under identical conditions in relation to the larval density. As index for the vitality, the mean number of attacked larch buds per larva could be used. Gradological stage (fig. 1) and, probably, larval density influence the vitality and the behaviour of the larvae (tabl. 3). In the present experiments, it could be observed that the rate of Eulophid parasitism increased with host density up to a certain level (tabl. 4). The limited searching capacity of the females probably rapidly limits the development of the importance of that parasites by decreasing host density.

     

Isolation and identification of a symbiotic bacterium fromSteinernema carpocapsae

Xenorhabdus nematophilus sp., an insect-pathogenic bacterium, was newly isolated from Korean entomopathogenic nematode ofSteinernema carpocapsae, which can be used as a useful bioinsecticide. Primary and secondary form variants ofXenorhabdus nematophilus were observed when culturedin vitro. Primary form variants adsorbed bromothymol blue, while secondary form did not. However, many other characters of two variants were very similar. The variants were all rod-shaped and cell size was highly variable ranging from 0.5 by 2.0 μm to 1.0 by 5.0 μm. Both produced highly toxic substances and killed the insect larva within 20–38 hr, indicating that insect pathogenicity ofXenorhabdus is not directly associated with its phase variation. In addition, cell-free culture supernatant ofXenorhabdus was sufficient to kill the insect larva by injecting it into insect hemolymph; however, cell-harboring culture broth was more effective for killing the insect. The use ofXenorhabdus nematophilus may provide a potential alternative toBacillus thuringiensis (Bt) toxins.     

Effect of parasitism byAscogaster reticulatus [Hym.: Braconidae] on growth of the host,Adoxophyes sp. [Lep.: Tortricidae]

The developmental interaction between the egg/larval parasitoid,Ascogaster reticulatus Watanabe (Hymenoptera: Braconidae) and its host,Adoxophyes sp. (Lepidoptera: Tortricidae) was examined. Prior to the egress of a final-instar parasitoid larva from the 4^th-instar host larva, host weight decreased by 22% from the maximum weight. The final body weight of a host larva was 27% of the maximum weight of a healthy 5^th-instar host. Food consumption was significantly reduced in both 3^rd-and 4^th-instar parasitized larvae compared with healthy ones. In the 4^th instar, a parasitized larva consumed 28% less artificial diet and produced less frass than a healthy larva. The growth rate of the endoparasitoid larvae greatly increased after their host's molt to the 4^th instar. Parasitoid larval volume increased 40 fold in the 4^th-instar host.      

Colour of larch heartwood and relationships to extractives and brown-rot decay resistance

Larch heartwood is appreciated for its good mechanical properties, its colour and its texture, and it is often used outdoors because of its natural durability (decay resistance). In this study the colour of larch heartwood was studied in relation to extractives and decay resistance, with the aim to estimate durability of larch heartwood from its colour. On a total of 293 trees colour in the CIE L*a*b* space (L* lightness, a* red/green axis, b* yellow/blue axis), extractives content (acetone and hot-water extractives, amount of phenolics) and the brown-rot decay resistance were determined. For calculating the relative decay resistance ( x), mass loss after inoculation for 16 weeks with two fungi [ Coniophora puteana (Schum.ex.Fr.) Karst., Poria placenta (Fr.) Cke, European standard EN 113] of larch heartwood samples was compared to Scots pine ( Pinus sylvestris L) sapwood reference samples (EN 350-1). Different species [Japanese larch ( Larix kaempferi Lamb.), Hybrid larch (Larix deciduax L. kaempferi) and European larch ( L. decidua Mill.)], provenances and age classes (38-year, >150-year) were included. Japanese larch heartwood turned out to be significantly more reddish (higher a*-values) compared to the European larch provenances. Reddishness of the hybrids was intermediate. The red hue (+a*) was strongly correlated with the amount of phenols ( r =0.84) and decay resistance ( r =0.63) and therefore suitable for prediction of both parameters. The results suggest that colour measurements of larch heartwood could be of benefit in tree breeding programs and for an optimised utilization of larch timber.     

Oviposition efficiency of the grey larch bud moth (Zeiraphera diniana) on different substrates and its relevance to the population dynamics of the moth

Females of the grey larch bud moth (Zeiraphera diniana Gn.) (Lepidoptera, Tortricidae) laid significantly more eggs on larch branches covered with naturally occurring Parmelia lichens than on branches without them.Numbers of eggs deposited and behaviour suggest that besides olfactory and mechanical stimuli from the antennae and the tarsi, thigmotactic stimuli perceived by the ovipositor play an important role for optimum oviposition.The fact that oviposition efficiency is reduced by 50% on larch branches from the lowland region of Europe where no lichens occur, is considered to be of importance for the determination of the fluctuations at low density in the lowland region.

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Zusammenfassung Die Weibchen des Grauen Lärchenwicklers (Zeiraphera diniana Gn.) legen auf Lärchenzweigen mit Parmelia-Flechten aus dem subalpinen Gradationsareal statistisch gesichert mehr Eier ab als auf Lärchenzweigen ohne Flechten aus dem collinen Verbreitungsgebiet.Die Anzahl und die Lage der abgelegten Eier sowie das Verhalten der Weibchen deuten darauf hin, dass thigmotaktische Reize des Ovipositors die vorausgegangene olfaktorische und mechanische Stimulation der Antennen und Tarsen ergänzen müssen, damit eine optimale Eiablage gewährleistet sei.Die Tatsache, dass die Ovipositionsleistung auf Lärchenzweigen ohne Flechten um ca. 50% verringert wird, dürfte von Bedeutung für die Determination des latenten Fluktuationstypus der Art sein.

     

Comparison of insect,fungal, and mechanically induced defoliation of larch: effects on plant productivity and subsequent host susceptibility

Summary Larch sawfly, Pristiphora erichsonii Hartig, and larch needlecast fungus, Mycosphaerella laricinia (R. Hartig) Neg., are early season defoliators restricted only to Larix host trees. Larch defoliation (100%) by either the fungus or insect, but not mechanical removal, induced systemic responses that reduced sawfly consumption and digestion rates one year later. In a feeding behavior assay, larvae quickly abandoned seedlings previously defoliated by M. laricinia. Adult female oviposition choice and egg deposition were unaffected. Seedling growth was not affected during the year of defoliation by M. laricinia, but was significantly reduced one year later. Defoliation by M. laricinia reduced stem volume, radial growth, root biomass and new shoot production. The latter tissue is the only oviposition resource for larch sawfly, and, in contrast, is not influenced by sawfly feeding. We hypothesize that M. laricinia infection may limit larch sawfly populations where both species coexist. However, this reduction is at a substantial net cost to larch productivity.     

Influence of Forest Types and Effects of Forestry Activities on Species Richness and Composition of Chrysomelidae in the Central Mountainous Region of Japan

Species richness and composition of the Chrysomelidae (Coleoptera) were studied in larch (Larix kaempheri [Lamb.] Carrière) plantations, secondary forests, and primary forests. In addition, the effects of forest management practices, such as thinning and long rotation, were examined in the larch plantation. The species richness of Chrysomelidae was higher in the larch plantation than in the secondary forest or in the primary forest. Among the larch plantations, the species richness in old-aged plantations was higher than that in middle-aged plantations. The composition of the beetle assemblages in the larch plantation differed from that in the secondary forest or in the primary forest. Exosoma akkoae (Chujo), Batophila acutangula Heikertinger, and Calomicrus nobyi Chujo were caught with a bias toward the larch plantation. Longitarsus succineus (Foudras) and Sphaeroderma tarsatum Baly were caught more in the secondary forest and the primary forest, respectively. More B. acutangula and S. tarsatum were caught in stands where their host plants occurred at higher rates. Species richness of understory plants was an important factor for chrysomlid species richness, and frequency of host occurrence affected the number of individuals of leaf beetles examined. It seems that forest types and forest management practices affect host plants as well as Chrysomelidae, and that these effects on the host plants also influence chrysomelid assemblages.     

Allopatric populations of the invasive larch casebearer differ in cold tolerance and phenology

1. Traits of non-native insect herbivores may vary spatially due to local genetic differences, rapid post-introduction evolution, and/or novel host plant associations. 2. Populations of larch casebearer, Coleophora laricella Hübner, originally from Europe have likely been isolated for > 60 years in North America on eastern larch, Larix laricina (Du Roi) K. Koch, and western larch, Larix occidentalis Nutt. 3. This study investigated cold tolerance and phenology of larvae collected from eastern larch in Minnesota, and western larch in Oregon, Idaho, and Montana, U.S.A. 4. Mean supercooling points of larvae from Minnesota were up to 10 °C lower than supercooling points of larvae from Oregon, Idaho, and Montana. 5. At ambient environmental conditions in spring, overwintering larvae from Minnesota required a mean (± SE) of 172 ± 19 degree-days above 5 °C to break winter quiescence and actively wander, significantly more than required by larvae from Oregon (66 ± 4), Idaho (64 ± 1), and Montana (60 ± 2). 6. Across all assays and despite substantial latitudinal and elevational variation among western larch sites, no significant differences in any traits were detected among larvae collected from western larch. 7. Spatial variation in cold tolerance and phenological traits of larch casebearer may be attributable to insect genetic differences and/or host plant effects, but exact mechanisms remain unknown. Differences in thermal biology between regions may result in disparate effects of climate change on insect populations and should be accounted for when forecasting insect dynamics across large spatial scales.     

Entomopathogenic symbiotic bacteria,Xenorhabdus and Photorhabdus of nematodes

Xenorhabdus and Photorhabdus species are entomopathogenic bacteria with a wide insect host range, that belong to the family Enterobacteriaceae. Xenorhabdus and Photorhabdus species symbiotically associate with nematodes of the families Steinernematidae and Heterorhabditidae respectively. The factor(s) determining the symbiotic interaction between nematodes and bacteria are yet to be identified. Xenorhabdus and Photorhabdus species exist in two main phenotypic forms, a phenomenon known as phase variation. The phase I (or primary form) varies from phase II (or secondary form) in certain physiological and morphological characteristics. There is no variation in the DNA integrity of phase I and phase II and this supports epigenetic regulatory mechanism in phase variation. Certain pathogenic determinants such as pili, lipopolysaccharides and toxins contribute to the pathogenicity of Xenorhabdus and Photorhabdus species, and both appear to be equally pathogenic to insects. The observed similarity in their virulence to insect hosts may reflect possible in vivo conversion of phase II to phase I, however the host cellular invasion and virulence is yet to be properly understood. The virulence of Xenorhabdus variants varies among insects apparently due to factors which include the feeding habits of the insects. The molecular mechanism and biological significance of phase variation are presently unknown.     

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.     

The influence of two endoparasitic wasps, <Emphasis Type="Italic">Hyposoter didymator</Emphasis> and <Emphasis Type="Italic">Chelonus inanitus</Emphasis>, on the growth and food consumption of their host larva <Emphasis Type="Italic">Spodoptera littoralis</Emphasis>

The influence of parasitism by Hyposoter didymator (Thunberg; Hymenoptera: Ichneumonidae) and Chelonus inanitus (Linnaeus) (Hymenoptera: Braconidae) on the growth and food consumption of their host Spodoptera littoralis (Boisduval) (Lepidoptera: Noctuidae) was studied in the laboratory. Parasitised larvae consumed significantly less artificial diet than unparasitised ones. Egg parasitisation by C. inanitus affected host larval consumption from the second day after emergence and it was significantly different from that of unparasitised ones. H. didymator, however, started to reduce larval consumption 4 days after parasitisation on the third instar host larvae. The overall reduction achieved by the larval endoparasitoid H. didymator is higher than that caused by the egg-larval endoparasitoid C. inanitus. The final body weight of a parasitised host larva by H. didymator and C. inanitus was only 6.7 and 13.0% of the maximum weight of an unparasitised sixth instar larva respectively. Moreover, parasitised larvae never reached the last instar. Results indicated that parasitised larvae might cause considerable less damage to the host plant than unparasitised ones.     

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.     

Summer temperature dependency of larch budmoth outbreaks revealed by Alpine tree-ring isotope chronologies

Larch budmoth (LBM, Zeiraphera diniana Gn.) outbreaks cause discernable physical alteration of cell growth in tree rings of host subalpine larch (Larix decidua Mill.) in the European Alps. However, it is not clear if these outbreaks also impact isotopic signatures in tree-ring cellulose, thereby masking climatic signals. We compared LBM outbreak events in stable carbon and oxygen isotope chronologies of larch and their corresponding tree-ring widths from two high-elevation sites (1800–2200 m a.s.l.) in the Swiss Alps for the period AD 1900–2004 against isotope data obtained from non-host spruce (Picea abies). At each site, two age classes of tree individuals (150–250 and 450–550 years old) were sampled. Inclusion of the latter age class enabled one chronology to be extended back to AD 1650, and a comparison with long-term monthly resolved temperature data. Within the constraints of this local study, we found that: (1) isotopic ratios in tree rings of larch provide a strong and consistent climatic signal of temperature; (2) at all sites the isotope signatures were not disturbed by LBM outbreaks, as shown, for example, by exceptionally high significant correlations between non-host spruce and host larch chronologies; (3) below-average July to August temperatures and LBM defoliation events have been coupled for more than three centuries. Dampening of Alps-wide LBM cyclicity since the 1980s and the coincidence of recently absent cool summers in the European Alps reinforce the assumption of a strong coherence between summer temperatures and LBM defoliation events. Our results demonstrate that stable isotopes in tree-ring cellulose of larch are an excellent climate proxy enabling the analysis of climate-driven changes of LBM cycles in the long term.     

Extracellular electrical activity from the photoreceptors of midge

The ontogeny of photosensitivity has been studied in a holometabolous insect, the midgeChironomus ramosus. The life cycle of midges shifts from an aquatic environment to a non-aquatic environment. Extracellular electrical activity of photoreceptor organs was recorded at larval and adult stages. We found an increase in photosensitivity as the larva metamorphosed to the adult stage. This is the first report of changes in photosensitivity during the development of any insect described in an ecological context.