A brachypterous butterfly?

Butterflies of the genus Redonda Adams & Bernard (Lepidoptera: Nymphalidae: Satyrinae) are endemic to the Andes of Venezuela. They comprise a monophyletic group of five allopatric taxa, females of which show various degrees of wing reduction and ability to fly. The female of Redonda bordoni Viloria & Pyrcz sp. nov. appears to be brachypterous and incapable of sustained flight, a phenomenon previously unknown within the Rhopalocera.     

Just beautiful?! What determines butterfly species for nature conservation

Prioritization is crucial in nature conservation, as land and financial resources are limited. Selection procedures must follow objective criteria, and not primarily subjective aspects, such as charisma. In this study, we assessed the level of charisma for all European butterflies. Based on these data, we analysed the charisma values of the species listed on the annexes of the EU Habitats Directive and of the species being of conservation priority according to criteria derived by three objective criteria: Species ecological specialisation, distribution, and threat. The mean level of charisma was higher for species of the EU Habitats Directive than for species of conservation priority and for not-listed species. Five of the twenty most charismatic species were also listed on the EU Habitats Directive, but none occurred on the list of species being of conservation priority. A trait space analysis revealed remarkable differences between the different species assortments: The species listed on the EU Habitats Directive covered a large trait space and included many species with high charismatic value, but low ecological and biogeographical relevance, while species of high conservation priority covered a restricted trait space and did not overlap with charismatic species. According to our findings, the selection of species for nature conservation still follows a mix of being aesthetic combined with some ecological criteria.

     

Orientation behaviour in the Grayling butterfly: thermoregulation or crypsis?

ABSTRACT.

• 1 Body orientation and wing tilting in Hipparchia semele (L.) have been explained previously both by the need for shadow-minimization (crypsis hypothesis) and by the need to regulate body temperature (thermoregulation hypothesis).
• 2 This study tests the thermoregulation hypothesis by quantifying butterfly orientation in relation to sun position, using vector analysis, and correlating the data with ambient temperature and time of day.
• 3 During sunshine, Graylings tend to present a maximal wing area to the sun's rays when the ambient temperature is relatively low, and a minimal area when it is high ('sunning’and‘heat-avoiding’positions respectively).
• 4 On cool sunny days, butterflies adopt sunning positions throughout the day, whereas on warm sunny days they adopt sunning positions during the outlying periods and heat-avoiding positions in the middle period of the day. Correlations between wing exposure and both ambient temperature and time of day are statistically significant for warm sunny days.
• 5 During overcast weather, butterfly orientation is very variable and possibly random.
• 6 Analysis of sunny day data supports the thermoregulation hypothesis. This is discussed in relation to other known types of thermoregulatory behaviour observed in this species: stilting, shade-seeking, ground-hugging and shivering.
• 7 Conversely, the results do not support the crypsis hypothesis.

     

Can we predict butterfly diversity along an elevation gradient from space?

An important challenge in ecology is to predict patterns of biodiversity across eco‐geographical gradients. This is particularly relevant in areas that are inaccessible, but are of high research and conservation value, such as mountains. Potentially, remotely‐sensed vegetation indices derived from satellite images can help in predicting species diversity in vast and remote areas via their relationship with two of the major factors that are known to affect biodiversity: productivity and spatial heterogeneity in productivity. Here, we examined whether the Normalized Difference Vegetation Index (NDVI) can be used effectively to predict changes in butterfly richness, range size rarity and beta diversity along an elevation gradient. We examined the relationship between butterfly diversity and both the mean NDVI within elevation belts (a surrogate of productivity) and the variability in NDVI within and among elevation belts (surrogates for spatial heterogeneity in productivity). We calculated NDVI at three spatial extents, using a high spatial resolution QuickBird satellite image. We obtained data on butterfly richness, rarity and beta diversity by field sampling 100 m quadrats and transects between 500 and 2200 m in Mt Hermon, Israel. We found that the variability in NDVI, as measured both within and among adjacent elevation belts, was strongly and significantly correlated with butterfly richness. Butterfly range size rarity was strongly correlated with the mean and the standard deviation of NDVI within belts. In our system it appears that it is spatial heterogeneity in productivity rather than productivity per se that explained butterfly richness. These results suggest that remotely‐sensed data can provide a useful tool for assessing spatial patterns of butterfly richness in inaccessible areas. The results further indicate the importance of considering spatial heterogeneity in productivity along elevation gradients, which has no lesser importance than productivity in shaping richness and rarity, especially at the local scale.     

High butterfly beta diversity between Brazilian cerrado and cerrado–caatinga transition zones

Tropical dry forests are among the most diverse and threatened habitats in the world, yet they are rarely protected and remain poorly studied. In Brazil, dry forests are naturally fragmented and embedded within various biomes, thus making it important to assess biotic homogeneity among geographically separated forest fragments. We sampled 7732 individuals belonging to 48 species to quantify the diversity of fruit-feeding butterfly communities at four Brazilian dry forest sites, and found differences in community structure between northern and central sites. Species richness per plot was the same in both areas, but abundance per plot was higher in northern sites. Species composition differed between sites mostly due to species of Satyrinae. Additive partitioning showed that beta diversity corresponded to 70.1% of all diversity. Rather than species loss, beta diversity primarily represented species turnover that was potentially driven by differences in the surrounding habitats. Butterfly community composition and abundance were influenced by vegetation where abundance increased with tree density and basal area, and decreased with the average tree height. Butterfly species richness and abundance were higher in the wet season than in the dry season, and all species sampled in the dry season were present in the wet season. Differences in community structure across relatively short geographic distances in the same type of habitat highlight the importance of conserving tropical dry forest fragments to ensure the maintenance of butterfly diversity and, presumably, other insects.     

Timing of butterfly parasitization of a plant–ant–scale symbiosis

In the Southeast Asian tropics, Arhopala lycaenid butterflies feed on Macaranga ant-plants inhabited by Crematogaster (subgenus Decacrema) ants tending Coccus-scale insects. A recent phylogenetic study showed that (1) the plants and ants have been codiversifying for the past 20–16 million years (Myr), and that (2) the tripartite symbiosis was formed 9–7 Myr ago, when the scale insects became involved in the plant–ant mutualism. To determine when the lycaenids first parasitized the Macaranga tripartite symbiosis, we constructed a molecular phylogeny of the lycaenids that feed on Macaranga by using mitochondrial and nuclear DNA sequence data and estimated their divergence times based on the cytochrome oxidase I molecular clock. The minimum age of the lycaenids was estimated by the time-calibrated phylogeny to be 2.05 Myr, about one-tenth the age of the plant–ant association, suggesting that the lycaenids are latecomers that associated themselves with the pre-existing symbiosis of plant, ant, and scale insects.     

Thoughts of a travelling ecologist 12.The plight of the monarch butterfly

<正>Nature conservation needs people who care about nature,and the efforts to convince people to care about the fate of nature has extensively relied on selected charismatic species,the"conservation icons".One famous example is the great panda,Ailuropoda melanoleuca,well known as the World Wide Fund for Nature(WWF)logo.When it comes to invertebrates,the     

Pollen amino acids—an additional diet for a nectar feeding butterfly?

The increase of the amino acid concentration over different time intervals in artificial nectar (i.e., a sucrose solution) due to pollen contamination was investigated in four Californian plant species (Aesculus californica, Amsinckia lunaris, Brodiaea pulchella, Carduus pycnocephalus), which are important nectar resources for a Californian colony of the butterflyBattus philenor as well as for other insects. The increase of the amino acid concentration in the medium is different in all four species and seems to be determined by a variety of factors including permeability of the pollen grain wall and presence or absence of pores. The results suggest a passive diffusion process of the free pollen amino acids into the medium rather than an active release. Implications from the experiments forBattus philenor and for other nectar feeding pollinators are discussed. A possible complementary effect of free pollen and nectar amino acids is proposed for plant species in which pollen is likely to be knocked into nectar by their flower visitors. A possible evolutionary pathway from nectar feeding butterflies such asBattus philenor to the complex derived pollen feeding habit in theHeliconius butterflies is proposed.     

Limited dispersal by the rare scarce heath butterfly—potential consequences for population persistence

Dispersal between habitat patches may be important for the long-term persistence of populations. We conducted a mark–release–recapture study and analysed the dispersal pattern in the scarce heath butterfly inhabiting a network of suitable habitat patches using stepwise logistic regression (SLR) and the Virtual Migration (VM) model. We also analysed the influence of different types of matrices. We found that the majority of the recaptured butterflies remained within the patch where they were originally caught. However, dispersal between patches did occur and both the SLR analysis and the VM model indicated that the migration pattern was significantly associated with patch area and its level of isolation. The SLR model also showed that there was a positive association between immigration rate and tree density, supporting earlier observations that this species prefers semi-open habitat. We discuss the use of SLR versus the VM model to analyse recapture data in dispersal studies. This system is not at equilibrium, as a number of the most important patches in the network are continuously being lost due to afforestation and a number of populations are facing deterministic extinction. This increases the risk of a chain reaction of local extinctions, which may cause a collapse of the whole system.     

Latitudinal gradients in butterfly body sizes: is there a general pattern?

We tested for the existence of latitudinal gradients in the body sizes of butterflies in North America, Europe, Australia and the Afrotropics. We initially compared body sizes (measured as male forewing length) of all butterflies found in 5° latitudinal bands in each region, and then evaluated the relationship between body size and latitude statistically using the latitudinal midpoint of each species' distribution. Trends were examined for species in all butterfly families together and for each family separately. We found that gradients in body sizes were inconsistent in different geographical regions and butterfly families; in some cases species were larger towards the tropics, in some they were smaller, and in other cases there were no relationships. Most of the gradients, when they existed, reflected between-family effects arising from changes in the relative numbers of species in each family across regions. We conclude that general ecological explanations for geographical trends in butterfly body sizes are inappropriate, and gradients largely reflect historical patterns of speciation within and between taxa in each biogeographical realm. Thus, the robustness of body size gradients found in other insect groups should be confirmed in future studies by including more than one geographical region whenever possible.     

Adaptive wing asymmetry in males of the speckled wood butterfly (Pararge aegeria)?

We analysed asymmetry in the wings of the speckled wood butterfly (Pararge aegeria)by measuring area, length and width of fore- and hindwings. The type of asymmetry is fluctuating except for fore- and hindwing area, and forewing width in males, where asymmetry is directional. The amount of asymmetry (variance of the left wing minus the right wing) is less in males than in females. Within males asymmetry was directional and less in pale, predominantly territorial males than in melanic, predominantly non-territorial males. Asymmetry was negatively related to growth rate within females, but not within males. Females grew faster than males, but had higher asymmetry, whereas the more asymmetrical melanic males grew more slowly than pale males. The differences in the type and amount of asymmetry between the sexes and colour classes suggest a relationship with sex-specific flight patterns such as the territorial spiralling flight of males. We hypothesize that slightly asymmetrical males turn faster, and therefore are superior in territorial disputes over more symmetrical or extremely asymmetrical males. This implies that sexual selection via male–male competition influences the type and amount of asymmetry. The existence of more extremely asymmetrical individuals in females, and to a lesser extent in non-territorial males, may indicate that there are costs in reducing asymmetry.     

Spatial ecology of host–parasitoid interactions: a threatened butterfly and its specialised parasitoid

Habitat conservation for threatened temperate insect species is often guided by one of two paradigms: a metapopulation approach focusing on patch area, isolation and number; or a habitat approach focusing on maintaining high quality habitat for the focal species. Recent research has identified the additive and interacting importance of both approaches for maintaining populations of threatened butterflies. For specialised host-parasitoid interactions, understanding the consequences of habitat characteristics for the interacting species is important, because (1) specialised parasitoids are particularly vulnerable to the consequences of fragmentation, and (2) altered interaction frequencies resulting from changes to habitat management or the spatial configuration of habitat are likely to have consequences for host dynamics. The spatial ecology of Cotesia bignellii, a specialist parasitoid of the threatened butterfly Euphydryas aurinia, was investigated at two spatial scales: within habitat patches (at the scale of individual aggregations of larvae, or ‘webs’) and among habitat patches (the scale of local populations). Parasitism rates were investigated in relation to larval web size, vegetation sward height and host density. Within patches, the probability of a larval webs being parasitized increased significantly with increasing number of larvae in the web, and parasitism rates increased significantly with increasing web isolation. The proportion of webs parasitized was significantly and negatively correlated with cluster density. Among habitat patches the proportion of parasitized webs decreased as cluster density increased. Clusters with a high proportion of larval webs parasitized tended to have lower parasitism rates per larval web. These results support the call for relatively large and continuous habitat patches to maintain stable parasitoid and host populations. Conservation efforts directed towards maintenance of high host plant density could allow E. aurinia to reduce parasitism risk, while providing C. bignellii with sufficient larval webs to allow population persistence.     

Male-killer dynamics in the tropical butterfly,Acraea encedana （Lepidoptera： Nymphalidae）

Sex ratio distortion in the tropical butterfly Acraea encedana is caused by infection with a male-killing bacterium of the genus Wolbachia. Previous research on this species has reported extreme female bias, high bacterial prevalences, and full sex role reversal. In this paper, we provide an assessment for the dynamics of the male-killer, based on a survey for sex ratios and Wolbachia prevalences among wild populations of A. encedana in Uganda. The study reveals that Wolbachia infection showed considerable variation over both spatial and temporal scales.     

Variation in butterfly egg adhesion: adaptation to local host plant senescence characteristics?

Most butterflies that overwinter as diapausing eggs have evolved oviposition behaviours where egg placement is near or on perennating portions of their host plant. We describe an alternative strategy used by alpine populations of a Lycaenid butterfly species complex where eggs are not attached strongly to the leaf substrate and subsequently fall off the plant. Other populations occurring at lower elevations attach their eggs strongly to the plant. In the laboratory, we measured the amount of force required to detach eggs from the leaf substrate. Eggs of the non‐alpine populations required more than five times the force required to detach eggs of the alpine population. Field surveys of the host plant characteristics used by various populations suggest that easy detachment may be adaptive for alpine populations because, unlike the host plants used by other populations, nearly all of the preceding year's vegetative growth is blown away by strong winds during the winter months.     

Butterfly morphology in a molecular age – Does it still matter in butterfly systematics?

We review morphological characters considered important for understanding butterfly phylogeny and evolution in the light of recent large-scale molecular phylogenies of the group. A number of the most important morphological works from the past half century are reviewed and morphological character evolution is reassessed based on the most recent phylogenetic results. In particular, higher level butterfly morphology is evaluated based on a very recent study combining an elaborate morphological dataset with a similar molecular one. Special attention is also given to the families Papilionidae, Nymphalidae and Hesperiidae which have all seen morphological and molecular efforts come together in large, combined works in recent years. In all of the examined cases the synergistic effect of combining elaborate morphological datasets with ditto molecular clearly outweigh the merits of either data type analysed on its own (even for ‘genome size’ molecular datasets). It is evident that morphology, far from being obsolete or arcane, still has an immensely important role to play in butterfly (and insect) phylogenetics. Not least because understanding morphology is essential for understanding and evaluating the evolutionary scenarios phylogenetic trees are supposed to illustrate.     

Conserving Europe’s most endangered butterfly: the Macedonian Grayling (Pseudochazara cingovskii)

The Macedonian Grayling is listed as critically endangered in the recent IUCN Red List of European butterflies because of its extreme rarity and habitat loss due to quarrying. This categorisation was, however, based on rather limited knowledge on its actual distribution, population size and habitat requirements. In 2012, we conducted field surveys to acquire more information. We found the species at six new sites extending its known range of suitable habitat to just under 10 km². The daily population size was estimated using capture-mark-recapture method in the most densely populated part of the Pletvar pass site at more than 650 individuals. Adults proved to be extremely sedentary, not moving far even within the continuous habitat on the same slope. Oviposition was observed on dry plant material and in a rock crevice close to the potential larval host plant Festuca sp. Quarrying is confirmed to be the main threat to the habitat of the Macedonian Grayling with five out of seven populated sites containing active marble quarries. Due to the enlargement of the known area of occupancy, its threat status would now be estimated at endangered. Despite the restricted knowledge about its distribution and trends in the population size, the IUCN criteria proved to be applicable to determine the threat status of a rare and localized butterfly such as Pseudochazara cingovskii. Its original assessment of being called the most threatened butterfly in Europe resulted in immediate research project and subsequent actions that will undoubtedly help to conserve it in the future.     

Population studies and conservation of Jamaica’s endangered swallowtail butterfly Papilio (Pterourus) homerus

This is an 18-year study of the endangered Papilio (Pterourus) homerus, adding substantial information to our scanty knowledge of its ecology. The contraction of a once contiguous but narrow population on a single Caribbean island carries the serious threat of extinction. There are now two populations or probably metapopulations, effectively isolated from each other. The butterfly’s larvae feed on Hernandia catalpaefolia and H. jamaicensis, both endemic to Jamaica, and development takes ～84 days from egg to the emerged adult. Adult numbers fluctuate rapidly, with peaks in July/August each year. Egg distribution was studied at three spatial levels: the food item (leaf cluster), the patch (tree) and the habitat (each valley). Major causes of developmental mortality were Chrysonotomyia, a eulophid parasitoid of the eggs, and bacterial infection of the larvae and pupae. Critically, the mortality from this wasp was lower in undisturbed forest than in the area disturbed by agriculture, this finding having important consequences for conservation. Although there was no evidence of a decline in numbers over the last century, we believe this is an artefact due to collectors working only at the periphery of its distribution. Even assuming that its population densities have not changed, the contraction of its usable habitat implies a similar reduction in average numbers and the small populations are susceptible to disaster. The efforts of researchers, NGOs, and Government agencies have greatly increased the level of awareness, making the people in some key areas the ‘protectors of the species’.     

Does habitat fragmentation affect temperature-related life-history traits? A laboratory test with a woodland butterfly

Habitat fragmentation may change local climatic conditions leading to altered selection regimes for life-history traits in small ectotherms, including several insects. We investigated temperature-related performance in terms of fitness among populations of the woodland butterfly Pararge aegeria (L.) originating from populations of a closed, continuous woodland landscape versus populations of an open, highly fragmented agricultural landscape in central Belgium. Female fecundity and longevity were evaluated in a temperature-gradient experiment. As predicted, females of woodland landscape origin reached higher maximum daily fecundity and lifetime number of eggs than did agricultural landscape females at low ambient temperatures, but this reversed at high ambient temperature. Egg weight decreased with temperature, and eggs of woodland butterflies were smaller. Contrary to what is generally assumed, remaining thorax mass was a better predictor of lifetime reproductive output than was abdomen mass. Since we used the F2 generation from wild-caught females reared under common garden conditions, the observed effects are likely to rely on intrinsic, heritable variation. Our results suggest that differential selection regimes associated with different landscapes intervene by intraspecific variation in the response of a butterfly to variation in ambient temperature, and may thus be helpful when making predictions of future impacts on how wild populations respond to environmental conditions under a global change scenario, with increasing temperatures and fragmented landscapes.