Host plants and butterfly biology. Do host‐plant strategies drive butterfly status?

Abstract. 1. To determine whether rarity and decline is linked to organism ecology, associations have been examined between butterfly larval host‐plant competitive, stress‐tolerant, ruderal (C‐S‐R) strategies and butterfly biology. 2. Associations have been sought between mean C‐S‐R scores for larval host plants with butterfly life history, morphology and physiology variables, resource use, population attributes, geography, and conservation status. Comparisons are carried out across species and controlled for phylogenetic patterning. 3. Butterfly biology is linked to host‐plant strategies. An increasing tendency of a butterfly's host plants to a particular strategy biases that butterfly species to functionally linked life‐history attributes and resource breadth and type. In turn, population attributes and geography are significantly and substantially affected by host choice and the strategies of these host plants. 4. The greatest contrast is between butterfly species whose host plants are labelled C and R strategists and those whose host plants are labelled S strategists. Increasingly high host‐plant C and R strategy scores bias butterflies to rapid development, short early stages, multivoltinism, long flight periods, early seasonal emergence, higher mobility, polyphagy, wide resource availability and biotope occupancy, open, areally expansive, patchy population structures, denser distributions, wider geographical ranges, resistance to range retractions as well as to increasing rarity in the face of environmental changes. Increasing host‐plant S strategy scores have reversed tendencies, biasing those butterfly species to extended development times, fewer broods, short flight periods, smaller wing expanse and lower mobility, monophagy, restricted resource exploitation and biotope occupancy, closed, areally limited populations with typical metapopulation structures, sparse distributions, and limited geographical ranges, range retractions, and increased rarity. 5. Species with S strategy host plants are species vulnerable to current environmental changes and species of conservation concern.     

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.     

Missing the target? A critical view on butterfly conservation efforts on calcareous grasslands in south-western Germany

Butterflies are strongly declining on grassland habitats of Central Europe. Therefore, the success of conservation measures on high quality grassland habitats is controversially discussed. We compared the changes in butterfly diversity and community structure on six managed calcareous grasslands with eight unmanaged vineyard fallows. We obtained strong losses of species diversity and remarkable shifts of community compositions on both habitat types. However, the changes on vineyard fallows were only slightly more severe but more stochastic than on the calcareous grasslands. The shifts in community composition with respect to functional species traits were rather similar between the two different grassland types so that complex butterfly communities evolved into generalist-dominated ones. Connectivity was higher among vineyard fallows than among calcareous grasslands. Consequently, conservation measures on calcareous grasslands only partly achieved their goal to maintain the high species diversity and functional complexity still observed in the 1970s. The negative impacts of eutrophication and monotonisation of the landscape as well as climate change are affecting all habitats, independently from management concepts. Therefore, management on conservation sites can buffer against these effects, but is not sufficient for a full compensation.     

Is the Natura 2000 network sufficient for conservation of butterfly diversity? A case study in Slovenia

Slovenia has one of the most extensive Natura 2000 networks in Europe with 259 SAC’s covering 31.4% of the country. To determine how well does the current network cover the areas of high butterfly diversity and/or aggregation of the butterfly species of conservation concern, the data from the recent survey for a distribution atlas were used. Altogether 99,423 records of 173 species collated after 1979 were used. The data distribution is slightly biased towards SAC’s, with 44.8% of localities within them, most likely due to sparsely sampled urban areas and intensive farmland areas which are found only outside SAC’s. The diversity and distribution of red listed species was evaluated at a 5 × 5 km grid square level. Additionally the importance of the size of the SAC’s was compared to their butterfly species diversity. In general the high diversity areas also hold the largest aggregation of red listed species with core areas concentrated in SW Slovenia. The SAC’s cover majority of areas with high diversity and the distribution of all but one threatened butterfly species. That species is Colias myrmidone, which is now considered extinct in Slovenia with no records after 1993. The most prominent areas with high conservation value in Slovenia not included in the SAC’s network are the Koro?ka region, Gori?ka Brda region, lower Sava River valley and Slovenske Gorice region. The butterfly diversity in small SAC’s is relatively high with increases in size only gradually increasing the species numbers, thus emphasizing the importance and conservation value of small SAC’s for sustaining high butterfly diversity in Slovenia.     

Do all inter-patch movements represent dispersal? A mixed kernel study of butterfly mobility in fragmented landscapes

1. In times of ongoing habitat fragmentation, the persistence of many species is determined by their dispersal abilities. Consequently, understanding the rules underlying movement between habitat patches is a key issue in conservation ecology. 2. We have analysed mark-release-recapture (MRR) data on inter-patches movements of the Dusky Large Blue butterfly Maculinea nausithous in a fragmented landscape in northern Bavaria, Germany. The aim of the analysis was to quantify distance dependence of dispersal as well as to evaluate the effect of target patch area on immigration probability. For statistical evaluation, we apply a 'reduced version' of the virtual migration model (VM), only fitting parameters for dispersal distance and immigration. In contrast to other analyses, we fit a mixed dispersal kernel to the MRR data. 3. A large fraction of recaptures happened in other habitat patches than those where individuals were initially caught. Further, we found significant evidence for the presence of a mixed dispersal kernel. The results indicate that individuals follow different strategies in their movements. Most movements are performed over small distances, nonetheless involving travelling between nearby habitat patches (median distance c. 480 m). A small fraction (c. 0·025) of the population has a tendency to move over larger distances (median distance c. 3800 m). Further, immigration was positively affected by patch area (I～A(ζ) ), with the scaling parameter ζ = 0·5. 4. Our findings should help to resolve the long-lasting dispute over the suitability of the negative exponential function vs. inverse-power one for modelling dispersal. Previous studies on various organisms found that the former typically gives better overall fit to empirical distance distributions, but that the latter better represents long-distance movement probabilities. As long-distance movements are more important for landscape-level effects and thus, e.g. for conservation-oriented analyses like PVAs, fitting inverse-power kernels has often been preferred. 5. We conclude that the above discrepancy may simply stem from the fact that recorded inter-patch movements are an outcome of two different processes: daily routine movements and genuine dispersal. Consequently, applying mixed dispersal kernels to disentangle the two processes is recommended.     

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.     

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.     

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.     

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.     

Do asynchronies in extinction debt affect the structure of trophic networks? A case study of antagonistic butterfly larvae–plant networks

Habitat loss and fragmentation affect species richness in fragmented habitats and can lead to immediate or time‐delayed species extinctions. Asynchronies in extinction and extinction debt between interacting species may have severe effects on ecological networks. However, these effects remain largely unknown. We evaluated the effects of habitat patch and landscape changes on antagonistic butterfly larvae–plant trophic networks in Mediterranean grasslands in which previous studies had shown the existence of extinction debt in plants but not in butterflies. We sampled current species richness of habitat‐specialist and generalist butterflies and vascular plants in 26 grasslands. We assessed the direct effects of historical and current patch and landscape characteristics on species richness and on butterfly larvae–plant trophic network metrics and robustness. Although positive species‐ and interactions–area relationships were found in all networks, structure and robustness was only affected by patch and landscape changes in networks involving the subset of butterfly specialists. Larger patches had more species (butterflies and host plants) and interactions but also more compartments, which decreased network connectance but increased network stability. Moreover, most likely due to the rescue effect, patch connectivity increased host‐plant species (but not butterfly) richness and total links, and network robustness in specialist networks. On the other hand, patch area loss decreased robustness in specialist butterfly larvae–plant networks and made them more prone to collapse against host plant extinctions. Finally, in all butterfly larvae–plant networks we also detected a past patch and landscape effect on network asymmetry, which indicates that there were different extinction rates and extinction debts for butterflies and host plants. We conclude that asynchronies in extinction and extinction debt in butterfly–plant networks provoked by patch and landscape changes caused changes in species richness and network links in all networks, as well as changes in network structure and robustness in specialist networks.     

Thermoregulatory differences between phenotypes in the speckled wood butterfly: hot perchers and cold patrollers?

Males of the speckled wood butterfly Pararge aegeria L. (Satyrinae), actively search for females (“patrolling”) or wait for them at particular places (“perching”). Darker males are more likely to patrol than pale ones, which are mainly territorial perchers. We studied whether this morphological variation relates to thermoregulatory differences. The relationship between thoracic temperature and ambient temperature differed between the colour types under natural conditions: darker males had on average lower body temperatures than paler males. Different activities (e.g. resting, flying) and behavioural strategies (perching or patrolling) were associated with differences in thoracic temperature: patrolling males which mainly engaged in long flights and periods of basking afterwards, had lower thoracic temperatures than perching males which engaged in very short flights, fights and basking. When resting for a while thoracic temperatures did not differ between males practising different strategies. Under laboratory conditions, darker males heated up faster than pale males but there was no difference in the thoracic temperature at which they started to fly. These results indicate that thermal requirements (or general conditions) differ between the behavioural strategies, and that behavioural differences between phenotypes (colour types) relate to differences in thermal ecology. This supports the idea that darker males are better adapted to patrolling. There is no evidence that one mate-locating strategy is always superior to the other, which coincides with the observation that both strategies co-exist. More generally, this study shows that relatively small differences in colour can have a considerable effect on thermoregulation and hence on the behavioural strategies a heliothermic insect will adopt. Received: 15 August 1997 / Accepted: 15 December 1997     

Does the presence of ant nests matter for oviposition to a specialized myrmecophilous Maculinea butterfly?

More than 50% of the lycaenid butterflies have an ant-associated lifestyle (myrmecophily) which may vary from coexistence to specific mutualistic or even parasitic interactions. Ant-related host-plant selection and oviposition has been observed in some myrmecophilous lycaenids. Therefore, it is remarkable that there is no evidence for this behaviour in the highly specialized, obligate myrmecophilous butterflies of the genus Maculinea. In contrast with previous findings, our results provide evidence for ant-related oviposition patterns in Maculinea alcon in relation to the distribution of specific host-ant nests (i.e. Myrmica ruginodis) based on repeated egg counts during the flight period in two populations. We also show that ant-related oviposition can be counterbalanced by intraspecific competition and oviposition deterrency when host plants already carry several eggs. Therefore, the absence of a correlation between egg load and the presence of host-ant nests at the end of the flight period should be interpreted carefully Whether ovipositional cues are obtained either directly (from ants or their nests) or indirectly (from vegetation structure), and whether alternative explanations based on the phenology and growth form of host plants are possible, is discussed.     

Biased correlated random walk and foray loop: which movement hypothesis drives a butterfly metapopulation?

Animals in fragmented landscapes have a major challenge to move between high-quality habitat patches through lower-quality matrix. Two current mechanistic hypotheses that describe the movement used by animals outside of their preferred patches (e.g., high-quality habitat or home range) are the biased, correlated random walk (BCRW) and the foray loop (FL). There is also a variant of FL with directed movement (FL_dm). While these have been most extensively tested on butterflies, they have never been tested simultaneously with data across a whole metapopulation and over multiple generations, two key scales for population dynamics. Using the pattern-oriented approach, we compare support for these competing hypotheses with a spatially explicit individual-based simulation model on an 11-year dataset that follows 12 patches of the federally endangered Fender’s blue butterfly (Plebejus icarioides fenderi) in Oregon’s Willamette Valley. BCRW and medium-scale FL and FL_dm scenarios predicted the annual total metapopulation size for ≥9 of 12 patches as well as patch extinctions. The key difference, however, was that the FL scenarios predicted patch colonizations and persistence poorly, failing to adequately capture movement dynamics; BCRW and one FL_dm scenario predicted the observed patch colonization and persistence with reasonable probabilities. This one FL_dm scenario, however, had larger prediction intervals. BCRW, the biologically simplest and thus most parsimonious movement hypothesis, performed consistently well across all nine different tests, resulting in the highest quality metapopulation predictions for butterfly conservation.