Pollination biology of a night‐flowering Galápagos endemic,Ipomoea habeliana (Convolvulaceae)

Ipomoea habeliana is an endemic, night‐flowering member of the Galápagos flora. Pollination experiments, flower‐visitor observations, nectar sampling, pollen transfer, and pollen to ovule ratio and pollen size studies were included in this project. The large, white flowers of this species set fruit via open pollination (55%), autonomous autogamy (51%), facilitated autogamy (91%), cross‐pollination (80%), diurnal open pollination (60%) and nocturnal open pollination (60%). Fruit set is pollen‐limited. Ants, beetles, crickets and hawk moths regularly visit the flowers. Ants are the most frequent visitors, but hawk moths are the only effective pollinators. Nectar is available throughout the night, but is most abundant early in the evening when hawk moth visits are most frequent. Experiments with fluorescent dust demonstrate intra‐ and inter‐plant pollen movement by hawk moths. Although this species is adapted for hawk moth pollination, it readily sets fruit via autonomous autogamy when no visits are made. Thus, it is concluded that it is facultatively xenogamous. Additional support for this conclusion is provided by the pollen to ovule ratio of 1407 and by the fact that the plants grow in a region that has few or no faithful pollinators. Conservation efforts for I. habeliana should include hand pollinations, which could significantly increase seed set. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 11–20.     

Extinction of the acoustic startle response in moths endemic to a bat-free habitat

Most moths use ears solely to detect the echolocation calls of hunting, insectivorous bats and evoke evasive flight manoeuvres. This singularity of purpose predicts that this sensoribehavioural network will regress if the selective force that originally maintained it is removed. We tested this with noctuid moths from the islands of Tahiti and Moorea, sites where bats have never existed and where an earlier study demonstrated that the ears of endemic species resemble those of adventives although partially reduced in sensitivity. To determine if these moths still express the anti-bat defensive behaviour of acoustic startle response (ASR) we compared the nocturnal flight times of six endemic to six adventive species in the presence and absence of artificial bat echolocation sounds. Whereas all of the adventive species reduced their flight times when exposed to ultrasound, only one of the six endemic species did so. These differences were significant when tested using a phylogenetically based pairwise comparison and when comparing effect sizes. We conclude that the absence of bats in this habitat has caused the neural circuitry that normally controls the ASR behaviour in bat-exposed moths to become decoupled from the functionally vestigial ears of endemic Tahitian moths.     

Neural evolution in the bat-free habitat of Tahiti: partial regression in an anti-predator auditory system

Noctuid moths endemic to the mountains of Tahiti have evolved in an environment without bats and these insects have lost a defensive behaviour against these predators, the acoustic startle response (ASR). The ASR in noctuid moths is presumed to be activated by a single auditory receptor neuron (A2 cell) and we report that while this cell still exists in endemic species and possesses similar sensitivity thresholds compared to the A2 cell of recently introduced species, it exhibits reduced firing activity to ASR-evoking sounds. This partial neural regression suggests that the evolutionary disappearance of the ASR in these insects is incomplete and that sensoribehavioural integration decays gradually following the removal of stabilizing selective forces.     

Visual position stabilization in the hummingbird hawk moth, Macroglossum stellatarum L. I. Behavioural analysis

Optomotor responses of freely flying hawk moths, Macroglossum stellatarum, were characterized while the animals were hovering in front of and feeding on a dummy flower. Compensatory translational and rotational movements of the hawk moth were elicited by vertical grating patterns moving horizontally, mimicking imposed rotational and translational displacements of the animal in the horizontal plane. Oscillatory translational and rotational pattern motion leads to compensatory responses that peak in the frequency range between 2 Hz and 4 Hz. The control systems mediating the translational and rotational components of the optomotor response do not seem to influence each other. The system mediating translational responses is more sensitive in the fronto-lateral part of the visual field than in the lateral part; the opposite is true for the rotational system. The sensitivity of the translational system does not change along the vertical, whereas the rotational system is much more sensitive to motion in the dorsal than in the ventral part of the visual field. These sensitivity gradients may reflect an adaptation to the specific requirements of position stabilization in front of flowers during feeding. Accepted: 13 August 1997     

Pollination of Platanthera chlorantha (Orchidaceae): new video registration of a hawkmoth (Sphingidae)

Documenting species interactions is a time consuming enterprise, in particular for rare interaction events and interactions taking place at night. Pollinators foraging on orchids have traditionally been monitored by discovering pollen vectors on collected insects, recording traces left by moths on the orchid, direct observations and recently by continuous video monitoring. Direct observations in the wild of orchids with low visitation rates is time demanding. In the present study I monitored greater butterfly‐orchids (Platanthera chlorantha), by using a event triggered video monitoring system. A total of 23 nights of monitoring were conducted, whereas only 6 nights had visits by one moth species, namely the pine hawk‐moth (Hyloicus pinastri). The total numbers of pine hawk‐moths registered were 18. In addition to species identification, the video recordings also enabled detection of pollinaria on the pine hawk‐moths. Most of the pine hawk‐moth visits took place around midnight. The visit lasted on average for 38.0 sec and the average number of flowers visited was 9.6. In future studies, this video system could give more details on interactions between orchids and insects and even link it to environmental factors (e.g. varying weather conditions).     

武夷山自然保护区天蛾科昆虫多样性的初步研究

通过2002年至2008年的陆续采集和2009年5月至10月定期系统调查采集,获得武夷山自然保护区的天蛾标本共2 103号,它们隶属于5亚科、33属、75种和亚种.该保护区天蛾科昆虫的区系成分以东洋区种类为主,其次是广布种,并且有少量的古北区种类及特有种;天蛾种类和个体数量的高峰期基本集中在6～8月,鹰翅天蛾Ambul...     

Nocturnal activity positively correlated with auditory sensitivity in noctuoid moths

We investigated the relationship between predator detection threshold and antipredator behaviour in noctuoid moths. Moths with ears sensitive to the echolocation calls of insectivorous bats use avoidance manoeuvres in flight to evade these predators. Earless moths generally fly less than eared species as a primary defence against predation by bats. For eared moths, however, there is interspecific variation in auditory sensitivity. At the species level, and when controlling for shared evolutionary history, nocturnal flight time and auditory sensitivity were positively correlated in moths, a relationship that most likely reflects selection pressure from aerial-hawking bats. We suggest that species-specific differences in the detection of predator cues are important but often overlooked factors in the evolution and maintenance of antipredator behaviour.     

Flight duration and flight muscle ultrastructure of unfed hawk moths

Flight muscle breakdown has been reported for many orders of insects, but the basis of this breakdown in insects with lifelong dependence on flight is less clear. Lepidopterans show such muscle changes across their lifespans, yet how this change affects the ability of these insects to complete their life cycles is not well documented. We investigated the changes in muscle function and ultrastructure of unfed aging adult hawk moths (Manduca sexta). Flight duration was examined in young, middle-aged, and advanced-aged unfed moths. After measurement of flight duration, the main flight muscle (dorsolongitudinal muscle) was collected and histologically prepared for transmission electron microscopy to compare several measurements of muscle ultrastructure among moths of different ages. Muscle function assays revealed significant positive correlations between muscle ultrastructure and flight distance that were greatest in middle-aged moths and least in young moths. In addition, changes in flight muscle ultrastructure were detected across treatment groups. The number of mitochondria in muscle cells peaked in middle-aged moths. Many wild M. sexta do not feed as adults; thus, understanding the changes in flight capacity and muscle ultrastructure in unfed moths provides a more complete understanding of the ecophysiology and resource allocation strategies of this species.     

Predation risk and mating behavior: the responses of moths to bat-like ultrasound

In the presence of predators, animals may reduce or alter theirmating activities. There has been little experimental studyof whether mating behavior varies with the level of predationrisk. Two species of moths, Pseudaletia unipuncta (Noctuidae)and Ostrinia nubilalis (Pyralidae), significantly reduced theirmate-seeking behavior under high levels of simulated predationrisk. Male moths aborted upwind flight in a pheromone plume,and females stopped releasing pheromone in response to soundssimulating the echolocation calls of bats. For O. nubilalis,but not for P. unipuncta, the response varied significantlywith the level of predation risk. Interspecific differencesin behavioral responses likely reflect differences in physiologicalauditory sensitivity and/or behavioral thresholds. Female behavioralresponses to sounds simulating the calls of bats that gleantheir prey from surfaces were significantly weaker than theirresponses to sounds resembling the calls of aerially hawkingbats; these results support the neurophysiological data thatthe calls of gleaning bats are relatively inaudible to moths.This study indicates that some animals can modify their reproductiveactivities in response to auditory cues from predators.     

The pollination of merremia palmeri (Convolvulaceae): can hawk moths be trusted?

The reproductive biology of Merremia palmeri and the pollination efficiency of its insect visitors were examined for a Sonoran Desert population in northwestern Mexico. Pollen transfer experiments proved that the plant is self-incompatible. Reproduction is, therefore, dependent upon reliable visitation by the primary pollinators, hawk moths. Many aspects of the floral structure are typical of sphingophilous flowers, and the time of flower opening and nectar secretion corresponded to the period of greatest hawk moth activity. A single hawk moth visit to a flower could be sufficient for successful fertilization. Additional visits up to five increased percentage fruit set, but flowers that received six or more visits had lower fruit and seed set. Neither the number of moth visits nor fruit and seed set were correlated with temperature or relative humidity. Over the course of the study >55% of flowers set fruit. We conclude that hawk moths are reliable and efficient pollinators for M. palmeri in a warm desert habitat.     

Frequency-intensity characteristics of cricket cercal interneurons: units with high-pass functions

Interneurons in the cercal sensory system of crickets respond in a cell-specific manner if the cercal hair sensilla are stimulated by air-particle oscillations at frequencies below about 2000 Hz. We investigated the filter properties of several of these interneurons, and tested the effect of stimulus intensity (typically 0.3–50 mm s⁻¹ peak-to-peak air-particle velocity) on the frequency response in the range 5–600 Hz. We focus on three interneurons (the lateral and medial giant interneurons and interneuron 9-3a) of Acheta domesticus which are characterized by a relatively high sensitivity above ca. 50–200 Hz. The responses of the medial giant interneuron usually increase monotonically with frequency and intensity. Interneuron 9-3a and the lateral giant interneuron exhibit saturation or response decrement at high frequencies and intensities. The lateral giant interneuron has an additional peak of sensitivity below about 40 Hz. Small individual variations in the relative locations of the two response areas of this interneuron within the frequency-intensity field are responsible for a large variability obtained if frequency-response curves are determined for particular intensities. Stimulus frequency does not affect the principal directional preferences of the three interneurons. Nevertheless, if tested individually, the lateral giant interneuron and interneuron 9-3a exhibit small changes of directional tuning. Accepted: 12 November 1997     

Acoustic irradiation produced by flying moths (Lepidoptera,Noctuidae)

Characteristics of acoustic waves accompanying the flight of noctuid moths (Noctuidae) were measured. The low-frequency part of the spectrum is formed of a series of up to 17 harmonics of the wingbeat frequency (30–50 Hz) with a general tendency toward the decrease in the spectral density and the increase in the sound frequency. The root-mean-square level of the sound pressure from flapping wings was found to be 70–78 dB SPL. Besides low-frequency components, the flight of moths was accompanied by short ultrasonic pulses, which appeared with every wingbeat. Most of the spectral energy was concentrated within a range of 7–150 kHz with the main peaks at 60–110 kHz. The short-term pulses were divided into two or more subpulses with different spectra. The high-frequency pulses were produced at two phases of the wingbeat cycle: during the pronation of the wings at the highest point and at the beginning of their upward movement from the lowest point. In most of the specimens tested, the peak amplitude of sounds varied from 55 to 65 dB SPL at a distance of 6 cm from the insect body. However, in nine noctuid species, no high-frequency acoustic components were recorded. In these experiments, the acoustic flow from the flying moth within a frequency range of 2 to 20 kHz did not exceed the self-noise level of the microphone amplifier (RMS 18 dB SPL). Probable mechanisms of the high frequency acoustic emission during flight, the effect of these sounds on the auditory sensitivity of moths, and the possibility of their self-revealing to insectivorous bats are discussed. In addition, spectral characteristics of the moth echolocation clicks were more precisely determined within the higher frequency range (>100 kHz).     

Pollination biology of mass flowering terrestrial utricularia species (lentibulariaceae) in the Indian Western Ghats

The pollination biology of three mass flowering Utricularia species of the Indian Western Ghats, U. albocaerulea, U. purpurascens, and U. reticulata, was studied for the first time by extensive observation of flower visitors, pollination experiments, and nectar analyses. The ephemerality of the Utricularia habitats on lateritic plateaus, weather conditions adverse to insects, lack of observations of flower visitors to other Utricularia spp., and the predominance of at least facultative autogamy in the few Utricularia species studied so far suggested that an autogamous breeding system is the common case in the genus. In contrast, we showed that the studied populations are incapable of autonomous selfing, or that it is an event of negligible rarity, although P/O was similarily low as in autogamous species investigated by other authors. In all three species the spatial arrangement of the reproductive organs makes an insect vector necessary for pollen transfer between and within flowers. However, U. purpurascens and U. reticulata are highly self-compatible, which allows for visitor-mediated auto-selfing and geitonogamy on inflorescence and clone level. Floral nectar is present in extremely small volumes in all three species, but sugar concentrations are high. More than 50 species of bees, butterflies, moths, hawk moths, and dipterans were observed to visit the flowers, and flower morphology facilitated pollination by all observed visitors. The results are discussed in the context of the phenological characteristics of the studied species, especially the phenomenon of mass flowering, and the environmental conditions of their habitats.     

Are butterflies and moths suitable ecological indicator systems for restoration measures of semi-natural calcareous grassland habitats?

The selection of suitable ecological indicator groups is of great importance for environmental assessments. To test and compare two such groups, we performed transect walks of butterflies and light traps of moths at eight sample localities in the Carinthian Alps. All of them were conducted with identical methods in the years 2002 and 2004 allowing the evaluation of the response on the conservation measures performed on five of the eight sites in late 2002. We recorded a total of 2346 butterflies (including Zygaenidae and Sesiidae) representing 83 species and 7025 moths of 534 species. 150 of these species were listed in the Red Data Book of Carinthia. In general, butterflies increased from 2002 to 2004 while moths declined. The highest increase rates of butterflies were obtained for the numbers of individuals of calcareous grassland specialists at the conservation sites, while their numbers were unchanged at the control sites. Similar trend differences between conservation and control sites were obtained for the Red Data Book butterfly species. On the contrary, the development of moth individuals was more positive at the control than the conservation sites for calcareous grassland specialists (only macro-moths) and species of the Red Data Book. However, change rates of species numbers were positively correlated between butterflies and moths. Principal Component Analysis revealed strong differences between the different sites, but mostly consistent results for butterflies and moths; however, stronger differences between years were only detected for some of the conservation sites for the butterfly communities. Our results show that butterflies as well as moths are suitable ecological indicator groups, but they do not yield identical results. Thus, butterflies are more suitable for the analysis of open habitats, whereas moths are suitable for open and forested habitats as well. Furthermore, butterflies might be a more sensitive indicator group than moths for the short-term detection of conservation measures, especially for the restoration of open habitat types.     

Morphological and physiological differences of the auditory system in three related bushcrickets (Orthoptera: Phaneropteridae, Poecilimon)

Abstract. The auditory system of three closely related bushcrickets was investigated with respect to morphological and physiological differences. The size of the acoustic vesicle in the prothorax cavity and the size of the acoustic spiracle were compared to differences in auditory tuning of the tympanic nerve and differences in the directionality. The results indicate that a small auditory vesicle and auditory spiracle provide reduced sensitivity in the high frequency range (above 10—15 kHz), but increase sensitivity at low frequencies (below 10 kHz). The directionality of the hearing system deteriorates at frequencies between 10 and 25 kHz in species with a small spiracle and trachea. The evolutionary implications of these differences of the auditory systems are discussed. They are considered to be influenced more by ecological factors than bioacoustical ones.     

Differential regulation of a homologous peptidergic neuron in grasshoppers: evolution of a neural circuit

The vasopressin-like immunoreactive (VPLI) neurons of grasshoppers have paired cell bodies in the suboesophageal ganglion and both anterior and posterior running axons. In non-oedipodine grasshopper species (e.g. Schistocerca gregaria), most of their arborisations are distributed in dorsal and lateral neuropil, while in oedipodine species (e.g. Locusta migratoria), the neurons have additional extensive axonal projections in both the optic lobes and proximal portions of the ganglionic peripheral nerves. This study demonstrates that these morphological differences correlate with their physiology. In L. migratoria, VPLI neuron activity is regulated primarily via a spontaneously active interneuron which descends from the brain. This descending interneuron is inhibited by a light-activated brain extraocular photoreceptor. Regulation of VPLI neuron activity by an extraocular photoreceptor is also seen in the other oedipodine grasshopper investigated. In the four non-oedipodines examined (from two subfamilies), we find no extraocular photoreceptor regulation of VPLI neuron activity. Despite this, VPLI neuron in S.␣gregaria does appear to be driven by a descending interneuron homologous to that in L. migratoria. The descending interneuron in both species receives similar mechanosensory input and excites the VPLI neuron via cholinergic synapses. Histamine injection into the medial protocerebrum of both species causes strong inhibition of the descending interneuron. The evolution of the neural circuitry, by which an extraocular photoreceptor comes to regulate the descending interneuron in oedipodine species, is discussed. Accepted: 6 January 1998     

Keeping up with bats: dynamic auditory tuning in a moth

Many night-flying insects evolved ultrasound sensitive ears in response to acoustic predation by echolocating bats . Noctuid moths are most sensitive to frequencies at 20-40 kHz , the lower range of bat ultrasound . This may disadvantage the moth because noctuid-hunting bats in particular echolocate at higher frequencies shortly before prey capture and thus improve their echolocation and reduce their acoustic conspicuousness . Yet, moth hearing is not simple; the ear's nonlinear dynamic response shifts its mechanical sensitivity up to high frequencies. Dependent on incident sound intensity, the moth's ear mechanically tunes up and anticipates the high frequencies used by hunting bats. Surprisingly, this tuning is hysteretic, keeping the ear tuned up for the bat's possible return. A mathematical model is constructed for predicting a linear relationship between the ear's mechanical stiffness and sound intensity. This nonlinear mechanical response is a parametric amplitude dependence that may constitute a feature common to other sensory systems. Adding another twist to the coevolutionary arms race between moths and bats, these results reveal unexpected sophistication in one of the simplest ears known and a novel perspective for interpreting bat echolocation calls.     

Colour vision: colouring the dark

Humans lose colour vision at night and it has often been assumed that this happens to other animals as well. It is not true of nocturnal moths, however: a recent study has shown that the elephant hawk moth makes use of trichromatic colour vision when seeking flowers by starlight.     

Responses of the less sensitive acoustic sense cells in the tympanic organs of some noctuid and geometrid moths

The separate impulses contributed by the A1 and A2 acoustic sense cells in the tympanic organs of the noctuids, Autographa pseudogamma and Noctua c.-nigrum, and by the A1, A2, and A3 sense cells in the tympanic organ of the geometrid, Ennomos magnarius, were identified and counted from oscillograms grams made as the moths were exposed to ultrasonic pulses of different intensities. These data were used to construct curves relating the response/intensity characteristics of the less sensitive acoustic sense cells to that of the most sensitive unit, A1. The A2 sense cells of the noctuids were found to be from 20 to 30 dB less sensitive than A1 at sound frequencies to which these ears are most sensitive. In the geometrid it was found that the A2 sense cell was 15 dB less sensitive than A1 and 12 dB more sensitive than A3. Only traces of the response of the fourth geometrid acoustic sense cell (A4) could be identified at high sound intensities. In both noctuids and geometrids the acoustic sensitivity of A2 relative to A1 remained unchanged when tested at selected ultrasonic frequencies between 28 and 50 kHz. This confirms the conclusion that the ears of these moths are incapable of pitch discrimination over this frequency range. Each of the systems had a dynamic range of 40 to 45 dB, that of the geometrid showing greater range overlap of the four A cells and hence greater capacity for sound intensity discrimination.     

Current understanding of the development of sex attractant-based biocontrol in burnet moths

Burnet moths, also known as the family Zygaenidae, are a typical diurnal family of Lepidoptera. Some species are important insect pests in agriculture and forestry. The use of sex attractants is one of the most important measures in the integrated pest management of burnet moths because these attractants are highly efficient, do not cause pollution, and are convenient and harmless to natural enemies. The earliest information about sex pheromones in Zygaenidae was reported in 1972. To date, many studies have described sex pheromones and attractants of species in Zygaenidae. Here, we review a total of 22 sex attractant compositions identified for 57 species of Zygaenidae and compare the differences in the structures of sex attractants among these species. The application of sex attractants in seasonal monitoring showed that the main activity period of zygaenid moths is from May to September each year. However, there are few reports on the effects of sex attractants on mass trapping of members of the family Zygaenidae. The ratio and degradation of sex attractants and the trap color, shape and hanging height were considered the main factors influencing capture effectiveness. Directions for further study of burnet moths are also discussed.