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Monitoring the response of butterfly communities to prescribed fire.

Federal land managers in the western United States are interested in the potential of prescribed fire as a tool to decrease fuel loads, increase vegetational heterogeneity, and increase faunal diversity in various ecosystems. I tested whether implementation of a prescribed fire program by the US Forest Service in a watershed in the central Great Basin had significant effects on butterfly species richness and composition. I monitored butterfly communities during the first two years after implementation in five to seven burn units and controls in the watershed. To estimate baseline spatial and temporal variation in butterfly communities in the greater ecosystem, I also monitored butterflies in five untreated canyons outside the project area. Butterfly species richness and butterfly species composition (measured as community similarity) did not differ significantly between burn units and controls. Geographic location had statistically significant effects on species richness. Butterfly species composition of individual locations varied over time, as did the magnitude of that variation. These results emphasize that standardized, repeatable monitoring protocols are vital for evaluating the effects of experimental management treatments and for predicting and assessing the effects of future management strategies and environmental changes.

Journal Article↗

Linking nectar amino acids to fitness in female butterflies.

Nectar of butterfly-pollinated flowers contains generally higher levels of amino acids than does nectar of flowers pollinated by most other animal types. One proposed explanation is that these amino acids promote butterfly fitness, although the evidence has been equivocal. In a new study, Mevi-Schütz and Erhardt showed that nectar amino acids enhanced fecundity in the butterfly Araschnia levana, but only when the larval diet was poor. Their results support the hypothesis that butterflies are agents of selection for higher nectar amino acid production, suggest that the larval food plant has a key role in the evolution of the flower-butterfly mutualism, and demonstrate that the importance, to butterfly reproduction, of different nutrient sources varies with butterfly nutritional state.

Journal Article↗

Double butterfly suture for high tension: a broadly anchored, horizontal, buried interrupted suture.

BACKGROUND: The excision of skin lesions such as tumors, nevi, and scars frequently results in tension on surgical wound margins. This tension is commonly counteracted surgically with buried, intracutaneous, interrupted sutures of absorbable material which are anchored vertically in the corium. METHOD: The horizontal, buried, intracutaneous butterfly suture has been described elsewhere. It is firmly anchored in the corium, everts wound margins, and adapts them nearly as broadly as two vertical sutures. It can also be laid as a double butterfly suture, as described here, and then has the shape of an "8." This double butterfly suture is equivalent to three vertical sutures because of its broad base in the corium. Moreover, it can cope with much greater tension because of its "pulley" effect. A single double butterfly suture usually suffices for small defects, particularly when the wound edges are cut obliquely with a longer rim of epidermis. MATERIALS: We have laid the traditional butterfly suture in more than 35,000 skin lesion excisions since 1985 and the double butterfly suture alone or as a supplement in more than 10,000 sutures since 1992. We use 2-0 to 6-0 polydioxanone for these procedures, since it has proven in trials to be the best-absorbed suture material. RESULTS: In most cases, the resulting scars were narrow and smooth in spite of high tension. Results were unsatisfactory in only 6.2% of procedures. CONCLUSION: The double butterfly suture described here has the advantages of withstanding tension better while everting wound margins and requiring fewer stitches for wound closure. However, it is important that the suture knot be deeply anchored beneath the corium.

Dermatologic Surgical Procedures↗

Role of photonic-crystal-type structures in the thermal regulation of a Lycaenid butterfly sister species pair.

One of the possible functions of the photonic-crystal structure found on the wing scales of some butterflies is investigated. The optical and electron microscopic investigation of two male butterflies-blue (colored) and brown (discolored)-representing a sister species pair and originating from different altitudes, revealed that the blue color can be attributed unambiguously to the fine, spongelike medium, called "pepper-pot structure," present between the ridges and the cross ribs in the scales of the colored butterfly. Only traces of this structure can be found on the scales of the discolored butterfly. Other physical measurements, mainly optical reflectivity, transmission, and thermal measurements, are correlated with structural data and simulation results. The thermal measurements reveal that under identical illumination conditions the high-altitude butterfly reaches a temperature 1.3-1.5 times the temperature reached by the low-altitude butterfly. This is attributed to the photonic-crystal-like behavior of the pepper-pot structure, which significantly reduces the penetration of light with wavelength in the blue region of the spectrum into the body of the scales. This sheds some light on the adaptation that enhances the survival chance of the butterfly in a cold environment rich in blue and UV radiation.

Journal Article↗

Behavioral foraging responses by the butterfly Heliconius melpomene to Lantana camara floral scent.

Floral color has been shown to influence flower selection by butterflies, but few studies have investigated the role of floral scent. In this study, adults of Heliconius melpomene L. (Lepidoptera: Nymphalidae: Heliconiinae) were tested in two-choice bioassays to investigate their ability to distinguish floral scent of the butterfly pollinated plant Lantana camara L. (Verbenaceae) from other plant scents. The relative importance of floral scent vs. color was also studied. Butterfly foraging behavior was measured as probing with proboscis. This probing, on floral models varying in scent and color, was timed. When given a choice of floral and vegetative scents of L. camara, newly emerged butterflies preferred floral scent, indicating an innate response to floral scents. When butterflies were conditioned to L. camara floral scent by offering the scent with yellow color and sugar water, yellow color elicited stronger feeding responses than did the floral scent. However, the floral scent of L. camara was preferred to that of the novel species Philadelphus coronarius L. (Hydrangiaceae). The floral scent of L. camara was dominated by tepenoid compounds, while that of P. coronarius by fatty acid derivatives, thus demonstrating totally different compositions. It is concluded that, while H. melpomene butterflies often use visual floral traits when selecting which flowers to visit, floral scents elicit behavioral responses that initiate and maintain foraging on flowers.

Animals↗

Virtual migration in tethered flying monarch butterflies reveals their orientation mechanisms.

A newly developed flight simulator allows monarch butterflies to fly actively for up to several hours in any horizontal direction while their fall migratory flight direction can be continuously recorded. From these data, long segments of virtual flight paths of tethered, flying, migratory monarch butterflies were reconstructed, and by advancing or retarding the butterflies' circadian clocks, we have shown that they possess a time-compensated sun compass. Control monarchs on local time fly approximately southwest, those 6-h time-advanced fly southeast, and 6-h time-delayed butterflies fly in northwesterly directions. Moreover, butterflies flown in the same apparatus under simulated overcast in natural magnetic fields were randomly oriented and did not change direction when magnetic fields were rotated. Therefore, these experiments do not provide any evidence that monarch butterflies use a magnetic compass during migration.

Animals↗

The aerodynamic costs of warning signals in palatable mimetic butterflies and their distasteful models.

Bates hypothesized that some butterfly species that are palatable gain protection from predation by appearing similar to distasteful butterflies. When undisturbed, distasteful butterflies fly slowly and in a straight line, and palatable Batesian mimics also adopt this nonchalant behaviour. When seized by predators, distasteful butterflies are defended by toxic or nauseous chemicals. Lacking chemical defences, Batesian mimics depend on flight to escape attacks. Here, I demonstrate that flight in warning-coloured mimetic butterflies and their distasteful models is more costly than in closely related non-mimetic butterflies. The increased cost is the result of differences in both wing shape and kinematics. Batesian mimics and their models slow the angular velocity of their wings to enhance the colour signal but at an aerodynamic cost. Moreover, the design for flight in Batesian mimics has an additional energetic cost over that of its models. The added cost may cause Batesian mimics to be rare, explaining a general pattern that Bates first observed.

Adaptation, Biological↗

Monitoring change in the abundance and distribution of insects using butterflies and other indicator groups.

Conservative estimates suggest that 50-90% of the existing insect species on Earth have still to be discovered, yet the named insects alone comprise more than half of all known species of organism. With such poor baseline knowledge, monitoring change in insect diversity poses a formidable challenge to scientists and most attempts to generalize involve large extrapolations from a few well-studied taxa. Butterflies are often the only group for which accurate measures of change can be obtained. Four schemes, used successfully to assess change in British butterflies, that are increasingly being applied across the world are described: Red Data Books (RDB) list the best judgements of experts of the conservation status of species in their field of expertise; mapping schemes plot the changing distributions of species at scales of 1-100 km2; transect monitoring schemes generate time series of changes in abundance in sample populations of species on fixed sites across the UK; and occasional surveys measure the number, boundaries and size of all populations of a (usually RDB) species at intervals of 10-30 years. All schemes describe consistent patterns of change, but if they are to be more generally useful, it is important to understand how well butterflies are representative of other taxa. Comparisons with similarly measured changes in native bird and plant species suggest that butterflies have declined more rapidly that these other groups in Britain; it should soon be possible to test whether this pattern exists elsewhere. It is also demonstrated that extinction rates in British butterflies are similar to those in a range of other insect groups over 100 years once recording bias is accounted for, although probably lower than in aquatic or parasitic taxa. It is concluded that butterflies represent adequate indicators of change for many terrestrial insect groups, but recommended that similar schemes be extended to other popular groups, especially dragonflies, bumblebees, hoverflies and ants. Given institutional backing, similar projects could be employed internationally and standardized. Finally, a range of schemes designed to monitor change in communities of aquatic macro-invertebrates is described. Although designed to use invertebrates as a bio-indicator of water quality for human use, these programmes could be extended to monitor the 2010 biodiversity targets of the World Summit on Sustainable Development.

Animals↗

Genetically engineered plants, endangered species, and risk: a temporal and spatial exposure assessment for Karner blue butterfly larvae and Bt maize pollen.

Genetically engineered maize (Zea mays) containing insecticidal endotoxin proteins from Bacillus thuringiensis (Bt) delta-endotoxin proteins has been adopted widely in the Midwestern United States. The proteins are toxic to several lepidopteran species and because a variety of maize tissues, including pollen, may express the endotoxins, the probability of exposure to nontarget species, including endangered species, needs to be understood. The objective of this study was to assess the potential temporal and spatial exposure of endangered Karner blue butterfly larvae (Lycaeides melissa samuelis) to Bt maize pollen in Wisconsin using probabilistic exposure techniques and geographic information systems analysis. Based on degree-day modeling of butterfly phenology and maize pollen shed, there is some potential for temporal exposure of larvae to maize pollen. However, in the majority of years and locations, maize pollen shed most likely will occur after the majority of larval feeding on wild lupine (Lupinus perennis). The spatial analysis indicates that some Karner blue butterfly populations occur in close proximity to maize fields, but in the vast majority of cases the butterfly's host plant and maize fields are separated by more than 500 m. A small number of potential or existing Karner blue butterfly sites are located near maize fields, including sites in two of the four counties where temporal overlap is most likely. The exposure assessment indicates that these two counties should receive the highest priority to determine if Karner blue butterfly larvae are actually at risk and then, if needed, to reduce or prevent exposure.

Animals↗

A wing expressed sequence tag resource for Bicyclus anynana butterflies, an evo-devo model.

BACKGROUND: Butterfly wing color patterns are a key model for integrating evolutionary developmental biology and the study of adaptive morphological evolution. Yet, despite the biological, economical and educational value of butterflies they are still relatively under-represented in terms of available genomic resources. Here, we describe an Expression Sequence Tag (EST) project for Bicyclus anynana that has identified the largest available collection to date of expressed genes for any butterfly. RESULTS: By targeting cDNAs from developing wings at the stages when pattern is specified, we biased gene discovery towards genes potentially involved in pattern formation. Assembly of 9,903 ESTs from a subtracted library allowed us to identify 4,251 genes of which 2,461 were annotated based on BLAST analyses against relevant gene collections. Gene prediction software identified 2,202 peptides, of which 215 longer than 100 amino acids had no homology to any known proteins and, thus, potentially represent novel or highly diverged butterfly genes. We combined gene and Single Nucleotide Polymorphism (SNP) identification by constructing cDNA libraries from pools of outbred individuals, and by sequencing clones from the 3' end to maximize alignment depth. Alignments of multi-member contigs allowed us to identify over 14,000 putative SNPs, with 316 genes having at least one high confidence double-hit SNP. We furthermore identified 320 microsatellites in transcribed genes that can potentially be used as genetic markers. CONCLUSION: Our project was designed to combine gene and sequence polymorphism discovery and has generated the largest gene collection available for any butterfly and many potential markers in expressed genes. These resources will be invaluable for exploring the potential of B. anynana in particular, and butterflies in general, as models in ecological, evolutionary, and developmental genetics.

Animals↗

Do monarch butterflies use polarized skylight for migratory orientation?

To test if migratory monarch butterflies use polarized light patterns as part of their time-compensated sun compass, we recorded their virtual flight paths in a flight simulator while the butterflies were exposed to patches of naturally polarized blue sky, artificial polarizers or a sunny sky. In addition, we tested butterflies with and without the polarized light detectors of their compound eye being occluded. The monarchs' orientation responses suggested that the butterflies did not use the polarized light patterns as a compass cue, nor did they exhibit a specific alignment response towards the axis of polarized light. When given direct view of the sun, migratory monarchs with their polarized light detectors painted out were still able to use their time-compensated compass: non-clockshifted butterflies, with their dorsal rim area occluded, oriented in their typical south-southwesterly migratory direction. Furthermore, they shifted their flight course clockwise by the predicted approximately 90 degrees after being advance clockshifted 6 h. We conclude that in migratory monarch butterflies, polarized light cues are not necessary for a time-compensated celestial compass to work and that the azimuthal position of the sun disc and/or the associated light-intensity and spectral gradients seem to be the migrants' major compass cue.

Animal Migration↗

Colour constancy in the swallowtail butterfly Papilio xuthus.

We have recently shown that the Japanese yellow swallowtail butterfly Papilio xuthus uses colour vision when searching for food. In the field, these butterflies feed on nectar provided by flowers of various colours not only in direct sunlight but also in shaded places and on cloudy days, suggesting that they have colour constancy. Here, we tested this hypothesis. We trained newly emerged Papilio xuthus to feed on sucrose solution on a paper patch of a certain colour under white illumination. The butterflies were then tested under both white and coloured illumination. Under white illumination, yellow- and red-trained butterflies selected the correctly coloured patch from a four-colour pattern and from a colour Mondrian collage. Under four different colours of illumination, we obtained results that were fundamentally similar to those under white illumination. Moreover, we performed critical tests using sets of two similar colours, which were also correctly discriminated by trained butterflies under coloured illumination. Taken together, we conclude that the butterfly Papilio xuthus exhibits some degree of colour constancy when searching for food.

Animals↗

Action spectrum of foraging behavior of the Japanese yellow swallowtail butterfly, Papilio xuthus.

This paper describes the action spectrum of foraging behavior of a butterfly, Papilio xuthus. We first established an experimental protocol to evaluate learning and discrimination of monochromatic light by the butterflies. We trained butterflies to feed on sucrose solution at the window illuminated with certain monochromatic light produced through a monochromator. After confirming that they learned the monochromatic light, after 10 days of training, we tested the butterflies one by one. We presented training wavelengths for each individual at different intensities, and recorded whether they perform foraging behavior under freely-flying as well as tethered conditions. Freely-flying butterflies responded to light by visiting the window and searching for nectar around it, whereas tethered butterflies responded by extending their proboscides towards the window. The light intensity required to elicit 50% response for each tested monochromatic light was plotted. The resulting action spectrum for the visit was rather flat with the maximum sensitivity a 420 nm, whereas the spectrum for the proboscis extension had prominent peaks at 380, 500 and 600 nm. The difference in action spectra indicates that the visit and the proboscis extension are controlled by two independent mechanisms at least in part.

Animals↗

Flavonoid sequestration by the common blue butterfly Polyommatus icarus: quantitative intraspecific variation in relation to larval hostplant, sex and body size.

Common blue butterflies (Polyommatus icarus) sequester flavonoids from their larval food and store these pigments as part of their adult wing colouration. Insects were reared on 10 different diets to assess effects of host plants on variation in flavonoid sequestration in this moderately polyphagous butterfly. Rearing experiments revealed an unexpectedly large gradient in flavonoid richness, ranging from individuals with high flavonoid loads (reared on inflorescences of Medicago sativa, Trifolium repens, T. pratense) to butterflies which contained almost no such pigments (fed with foliage of M. sativa or Robinia pseudoacacia). Flavonoid sequestration was much more effective from natural hostplants than from experimentally offered diets which would not be accepted in the field. Female butterflies on average sequestered almost 60% more flavonoids than males. This sex difference was more pronounced on natural than on experimental diets. Flavonoid load was significantly and positively related to dry mass and forewing length as two important fitness correlates of butterflies. This correlation was particularly strong on experimental diets (i.e. under constraining conditions for development). On natural hostplants, in contrast, when butterflies generally were flavonoid-rich, no clear relationship between flavonoid load and size or mass emerged. Our analytical data are consistent with field results according to which females rich in UV-absorbing flavonoid wing pigments are more attractive to mate-searching males. In P. icarus, flavonoid richness might therefore increase visibility (by more effective sensory stimulation of the visual system), but could also confer information about the feeding history, and thus ontogenetically determined 'quality' of a potential mate.

Journal Article↗

Do neotropical migrant butterflies navigate using a solar compass?

Many tropical butterfly species are well-known for their migratory behaviour. Although these insects can maintain a constant direction throughout the day, the physiological mechanisms of orientation are unknown. It has been argued that tropical migrant butterflies must use a time-compensated sun compass to accomplish their journey, but the crucial experimental manipulations to test this hypothesis have not been conducted. This study reports the results of clock-shift experiments performed with two species of migrating butterflies (Pieridae: Aphrissa statira and Phoebis argante) captured during flight across Lake Gatun, Panama. The observed constant flight bearing of natural controls suggests that these species are capable of performing time-compensated celestial navigation. Our clock-shift experiments suggest that a sun compass is involved. Individuals submitted to a 4 h advance shift took significantly different mean orientations on release compared with control butterflies. The direction of this difference was consistent with the use of a sun compass. The magnitude was approximately half the predicted value if the vanishing bearing of released butterflies was used as the variable to evaluate the effect of time-shifting and approximately three-quarters of that predicted if the estimated heading was the variable used. Mean vanishing bearings of control and experimental butterflies did not correspond to predicted values. This difference can be attributed largely to the combined effects of wind and handling.

Journal Article↗

In search of butterfly origins.

To determine the sister taxon to butterflies, the relationships among the macrolepidopteran superfamilies were investigated using sequence data from the ND1 gene of mitochondrial DNA. Both sequence data and translated amino acids were used. We examined how different models of amino acid evolution (nonadditive) affected the inference of tree topology. We then added previously published sequence data from the 18S and 28S subunits of ribosomal RNA. In most analyses, regardless of data type and treatment, the Hedyloidea is either the sister taxon to butterflies or derived within the butterfly clade. These molecular results are compared to a reanalysis of morphological characteristics. The reanalysis of morphology agrees with Minet's 1991 hypothesis of hedyloid relationships. Analysis of combined molecular and morphology data supports the hypothesis that hedylids are the sister group to butterflies. Scott's hypothesis that hawk moths (Sphingidae) are the sister taxon to butterflies is discounted.

Amino Acid Sequence↗

The effect of military training activity on eastern lupine and the Karner blue butterfly at Fort McCoy, Wisconsin, USA.

The US Department of Defense (DOD) manages over 10.1 million ha of land, much of which is used for training military personnel. However, vast sections receive little or no use, and military lands have become refuges for many species. At Fort McCoy, Wisconsin, USA, populations of the endangered Karner blue butterfly (Lycaeides melissa samuelis) are found in oak and pine barren communities where wild lupine (Lupinus perennis), a perennial forb required by Karner blue butterfly larvae, still occurs. Oak and pine barren communities are disturbance-dependent, and the barrens ecosystems in the Midwest have declined in extent by 98% because of fire suppression, succession, and habitat fragmentation. We studied the effects of disturbance by military manuever training on the density of lupine and Karner blue butterfly at Fort McCoy. We also wanted to determine whether military training activity could enhance Karner blue butterfly habitat. At locations where tracked vehicles had driven through lupine patches, the abundance of lupine and nectar-producing plants was greater in the median strip between vehicle ruts than in vehicle ruts or 5 m outside the vehicle ruts. The proportion of lupine stems with Karner blue butterfly larvae feeding sign (the ratio of stems fed upon to stems examined) was greater in areas where military vehicles had traveled than where they had not. The proportion of lupine stems with feeding sign and lupine stem density was also positively related to the occurrence of prior bivouacs and fires caused by military munitions. Shrub and forest canopy abundance were lower in areas traveled by tracked vehicles. At the scale of the lupine patch, lupine abundance and the proportion of lupine stems with feeding sign were positively correlated with military training activities, suggesting that maintenance of lupine habitat can be achieved in concert with military training.

Animals↗

Local species immigration, extinction, and turnover of butterflies in relation to habitat area and habitat isolation.

Temporal dynamics of insect communities in terrestrial habitat fragments have been rarely studied. Here it was tested whether immigration, extinction, and turnover of butterfly species change with area and isolation of 31 calcareous grasslands. The area ranged from 0.03 to 5.14 ha, the isolation index from 2,100 to 86,000 (edge-to-edge distance 55-1,894 m). In both study years (1996, 2000), the total number of individuals (16,466, 15,101) and species (60, 54) sampled across all sites were similar and number of species increased with area in both years indicating an equilibrium. Rates of extinction (38% for habitat specialists vs. 20% for generalists) and turnover (51% vs. 35%) were higher, and rates of immigration (11% vs. 30%) were lower for habitat specialists than for generalists. Extinction and turnover rates decreased with increasing fragment size for both specialist ( n=25 species) and generalist ( n=36) butterflies, but specialists showed a significantly steeper decrease with increasing fragment size than generalists. Immigration rates increased with area. As a result, species number of habitat specialists declined in small habitats but not in large habitats between 1996 and 2000. No significant impact of habitat isolation on the butterfly community was found. The data suggest that large habitat fragments are of special importance for the conservation of the specialized, most endangered butterfly species. Habitat isolation appears to be less important, as butterflies can cope with the habitat mosaic in our study region.

Animals↗