Phenology of weight, moisture and energy reserves of Australian monarch butterflies, Danaus plexippus

Abstract. 1. In the Sydney area of New South Wales, dry weights of reproductive monarch butterflies averaged 156 mg and were higher during winter than in other seasons. Dry weights of non-reproductive monarchs ranged from 216 to 324 mg and declined by 15–25% during over wintering.
2. Fat in reproductive butterflies ranged from 0.009 to 0.017g/0.1g dry weight and was lower during winter than in other seasons. Fat content of non-reproductive monarchs was higher (0.019–0.037 g/0.1g) and declined by 24–51% during over wintering.
3. Lean dry weights of reproductive monarchs were lower than those of non-reproductive individuals. Lean dry weights of non-reproductive butterflies increased rapidly at the beginning of over wintering and remained high throughout the winter. Analysis of protein content indicated the higher lean weight of non-reproductive monarchs was due to greater protein levels.
4. Moisture content of monarchs did not vary with season or reproductive status and appeared to be correlated with ambient humidity.
5. Non-reproductive monarch butterflies in New South Wales adjust bio chemically during over wintering. Energy reserve dynamics of these butterflies are comparable to those that occur in non-reproductive monarchs in North America.     

Preferences for nectar sugars in the peacock butterfly, Inachis io

1. Peacock butterflies, Inachis io , were tested experimentally for their preferences for nectar sugars.
2. In tests with different plain sugar solutions (25%, weight to total weight) the butterflies strongly preferred sucrose and fructose over glucose. They also preferred sucrose over fructose.
3. In tests with mixed sugar solutions the butterflies clearly preferred both sucrose-dominant (sucrose : hexoses = 5 : 1) and balanced sugar solutions (sucrose : glucose : fructose = 1 : 1 : 1) over hexose-dominant sugar solutions (sucrose : hexoses = 1 : 5).
4. Females consumed significantly more of the balanced sugar solution than did males.
5. These results are discussed with respect to previous experiments on nectar preferences of butterflies, nectar sugar composition of butterfly-pollinated flowers, and flower preferences, physiological and reproductive aspects of butterflies.     

Conservation of arthropod diversity in montane wetlands: effect of altitude, habitat quality and habitat fragmentation on butterflies and grasshoppers

1. The patterns of arthropod diversity were investigated in 24 montane wetlands in Switzerland. These differed in altitude, management regime (cattle-grazing vs. mowing), vegetation structure (index combining vegetation height and density) and degree of habitat fragmentation.
2. The general arthropod diversity was determined by net sampling at 10 sampling points per site. The diversity of grasshoppers and butterflies was measured by counting species richness at the site and species density (species richness per unit area) on transects. The species richness of grasshoppers and butterflies was found to be more sensitive to the geographical attributes of the site whereas species density was more affected by the habitat quality.
3. Grasshopper diversity decreased within the observed altitudinal range (800–1400 m) and was higher at grazed sites, whereas butterfly diversity was higher at mown sites. Arthropod diversity but not abundance of arthropods was positively related to the vegetation structure.
4. The species richness of butterflies was negatively influenced by the degree of habitat fragmentation: both the size of habitat as well as the area of wetland habitats within 4 km were related positively to the number of specialist wetland butterflies.
5. Late mowing as well as low-density cattle-grazing are appropriate management actions to maintain arthropod diversity in montane wetlands. In order to establish site-specific management plans, the biology of the present target species as well as the historical context should be considered.
6. We suggest that the best protection for the species examined in this study would be a network of wetland sites managed using a variety of traditional, non-intensive methods. This can only be achieved by coordinated planning of conservation measures among sites.     

Effects of maternal age and adult diet on egg weight in the butterfly Euphydryas editha

ABSTRACT. 1. Euphydryas editha (Boisduval) (Nymphalidae) butterflies exhibited no consistent pattern of change in egg weight over their lifetimes. This was the case for captive butterflies fed diets containing sugars and amino acids, for butterflies fed diets containing only sugar, and for those left completely at liberty in their natural habitats between ovipositions.
2. Two aspects of these results conflict with the findings of a previous study on the same subspecies. First, the previous study found that egg weight decreased with increasing maternal age. Second, this decrease was mitigated by addition of amino acids to the adult diet.
3. We accept the findings of these previous workers that egg weight decreased with maternal age under their laboratory conditions. The discrepancy between their data and ours is likely to stem from the more natural conditions under which our insects were kept. Our own data from captive insects are not distinguishable from our field data.
4. These results suggest that declines of egg weight with maternal age found in studies of other insects may also result from the artificial nature of laboratory conditions. We question whether such declines are general.
5. We suggest that the second discrepancy between the two sets of work (effects of amino acids) stems from a design problem in the previous study.     

An experimental examination of the effects of habitat quality on the dispersal and local abundance of the butterfly Parnassius smintheus

Abstract 1. Nectar flower abundance was manipulated through flower removal, and sex ratio was manipulated by moving individual butterflies within a series of nine alpine meadows. The movement and abundance of the butterfly Parnassius smintheus in the meadows were monitored using mark–release–recapture methods.
2. A total of 937 butterflies, 698 males and 239 females, was captured. There were 223 observed between-meadow movements. Fifty-two per cent of males and 35% of females moved among meadows.
3. The immigration of male butterflies was related positively to nectar flowers, host plant abundance, and female butterflies. Male emigration was not affected by any of the treatments. The number of males captured was related positively to nectar flowers and host plants but not affected by sex ratio. The number of resident male butterflies was greater in meadows containing flowers and was related positively to host plant abundance, but unaffected by sex ratio.
4. Flower removal, sex ratio, and abundance of Sedum had no significant effect on the abundance, movement, or residence time for female butterflies, in part due to small sample size.
5. The fact that males immigrate to higher quality meadows suggests that male butterflies are assessing meadow quality, either by sampling meadows or potentially from a distance using olfactory cues.     

Energy reserves and metabolic expenditures of monarch butterflies overwintering in southern California

Abstract. 1. Energetic expenditure and predicted requirements for overwintering metabolism were determined for monarch butterflies ( Danaus plexippus L.) in southern California.
2. Fat content of butterflies declined steadily from a maximum of 71% lean dry weight in late November to a minimum of 36% lean dry weight in late January. The energy expenditure measured by fat depletion over a 61 day period from 24 November to 25 January was 26.05 joules per day.
3. Butterflies were generally the same temperature as the air at any time they were inactive, whether they were part of a large cluster or roosting solitarily.
4. Oxygen consumption of butterflies increased in a curvilinear fashion with increasing air temperature. Thus, the predicted metabolic requirements for an inactive butterfly during their quiescent period from late November to late January was 30.32 joules per day.
5. In contrast to the steady depletion of fat reserves during their quiescent period in December and January, low and stable fat levels of butterflies in late February coincide with high levels of flight activity, mating and emigration of females from the colony.
6. Progressive tightening of the energy balance due to increasing nocturnal temperatures from northern to southern California coastal areas may limit the southern distribution and duration of overwintering aggregations.     

Climate-based models of spatial patterns of species richness in Egypt's butterfly and mammal fauna

Aim  Identifying areas of high species richness is an important goal of conservation biogeography. In this study we compared alternative methods for generating climate-based estimates of spatial patterns of butterfly and mammal species richness.
Location  Egypt.
Methods  Data on the occurrence of butterflies and mammals in Egypt were taken from an electronic database compiled from museum records and the literature. Using M axent , species distribution models were built with these data and with variables describing climate and habitat. Species richness predictions were made by summing distribution models for individual species and by modelling observed species richness directly using the same environmental variables.
Results  Estimates of species richness from both methods correlated positively with each other and with observed species richness. Protected areas had higher species richness (both predicted and actual) than unprotected areas.
Main conclusions  Our results suggest that climate-based models of species richness could provide a rapid method for selecting potential areas for protection and thus have important implications for biodiversity conservation.     

Parasitism by the mite Trombidium breei on four U.K. butterfly species

Abstract 1.  The incidence of parasitism by larvae of the mite species Trombidium breei was reported in one population of the lycaenid butterfly Polyommatus icarus , four populations of the satyrine butterfly Maniola jurtina , one population of the satyrine butterfly Aphantopus hyperanthus , and two populations of the satyrine butterfly Pyronia tithonus , as well as on one specimen of the dipteran Alophorus hemiptera . A considerable proportion of butterflies (11-50%) was infested in all study populations.
2. The pattern of infestation was examined in detail in M. jurtina . Males had a significantly higher incidence of infestation than females, and middle-aged butterflies had a higher incidence of infestation than old or young butterflies. The incidence of infestation peaked in the middle of the flight season, and this seasonal effect was independent of the effect of butterfly age.
3. Using a model based on capture-recapture data, it was estimated that a hypothetical ideal male M. jurtina that lives exactly the mean expected lifespan of 9-10 days has an approximately 75% chance of becoming infested with mites at least once during its lifetime, a mean time to first infestation of 3-4 days, and an average infestation persistence time of 2-3 days.
4. Capture-recapture data failed to show any effect of mite infestation on the lifespan or within-habitat movement rate of M. jurtina .
5. In experiments in which individual butterflies were taken out of their normal habitat and released, M. jurtina and P. tithonus that were infested with mite larvae did not differ from uninfested individuals in the efficiency with which they returned to suitable habitat. Thus, parasitism by T. breei larvae had no detectable effects on flight performance or orientation ability.
6. The results suggest that trombidiid mite larvae have limited potential in the biological control of insect pests.     

The case for headland set-aside: consideration of whole-farm gross margins and grain production on two farms with contrasting rotations

Spreadsheet calculations were used to compare headland set-aside with rotational set-aside in terms of gross margin and grain production on two farms with contrasting rotations. At Broom's Barn, Suffolk there was a five course rotation consisting of sugar beet and four cereals, while at Bunny Park, Nottinghamshire oilseed rape was the break crop, followed by three cereals. For both farms a sensitivity analysis was used to investigate the effect of the proportion of the farm required to be set aside, the extent of headland yield reduction and the cereal price on the outcome of the spreadsheet calculations. In general headland set-aside out-performed rotational set-aside. Yields on headlands were always less than the main body of the field, but it was on the turning headlands where yield was particularly depressed. Thus the advantage, in terms of gross margin, of setting aside the headlands was greatest when the requirement could be fulfilled by fallowing most turning headlands, but no non-turning headlands. Generally, rotational set-aside reduced grain production more than headland set-aside, but when the spreadsheet models were adjusted to maintain the area of sugar beet, the situation was reversed. In this case headland set-aside produced a larger gross margin and had the greatest impact on grain production. An extension of the analysis is to ask the question: even if set aside is not an option, is it worthwhile to crop rather than fallow headlands? For cereals and oilseed rape the answer is in the affirmative; the gross margin for headlands remains positive, even when prices are much reduced, because the arable area payment more than offsets the variable costs. This does not hold for crops such as sugar beet that are not eligible for arable area payments. An additional benefit from headland set-aside is its potential to enhance the environment through increased habitat diversity and the provision of ‘buffer zones’ to prevent agrochemicals from drifting into hedges and watercourses.     

Changes in egg colour, egg weight and oviposition rate with the number of eggs laid by wild females of the small heath butterfly, Coenonympha pamphilus

ABSTRACT. 1. The colour of eggs laid by individual Coenonympha pamphilus (L.) (Lep., Satyridae) females changed over their lifespan. The first laid eggs were green, but after having laid about 100 eggs, females laid only yellow eggs.
2. By following females in the field, or by capturing wild females, and noting the colour of their eggs, it was established that younger females showed higher oviposition rates and laid heavier eggs than older females.
3. Hence, wild C.pamphilus females are unable to sustain a constant egg production by extracting amino acids or any other nutrients from nectar. We hypothesize that this is a general phenomenon in nectar feeding butterflies.
4. The egg colour did not seem to match the colour of the substrate on which the female chose to deposit the egg.     

Species richness in agroecosystems: the effect of landscape, habitat and farm management

It has been suggested that biodiversity in agroecosystems depends on both landscape heterogeneity and farm management, but at the same time, studies of biodiversity in relation to both landscape variables and farm management are rare. We investigated the species richness of plants, butterflies, carabids, rove beetles and the diversity of spiders in cereal fields, leys (grass and clover crop) and semi-natural pastures at 16 farms in Central East Sweden. The farms were divided into eight pairs of one conventional and one organic farm to enable us to separate the effects of landscape and farm management on biodiversity. The pairing was based on land use, location, and landscape features. Species richness of different taxonomic groups was generally not correlated. There were no differences in species richness between the farming systems, except for carabids that had higher numbers of species on conventional farms. The species richness generally increased with landscape heterogeneity on a farm scale. Habitat type had a major effect on the species richness for most groups, with most species found in pastures and leys. The correlations between species richness and landscape variables on a farm scale, and not on a scale of multiple farms, identify farmers as the important decision-maker in conservation issues for these taxonomic groups. We discuss the role of species richness of pests' natural enemies for biological control and conservation strategies of the more common species in the agricultural landscape.     

Impacts of Climate-Change-Driven Sea Level Rise on Intertidal Rocky Reef Habitats Will Be Variable and Site Specific

Intertidal rocky reefs are complex and rich ecosystems that are vulnerable to even the smallest fluctuations in sea level. We modelled habitat loss associated with sea level rise for intertidal rocky reefs using GIS, high-resolution digital imagery, and LIDAR technology at fine-scale resolution (0.1 m per pixel). We used projected sea levels of +0.3 m, +0.5 m and +1.0 m above current Mean Low Tide Level (0.4 m). Habitat loss and changes were analysed for each scenario for five headlands in the Solitary Islands Marine Park (SIMP), Australia. The results indicate that changes to habitat extent will be variable across different shores and will not necessarily result in net loss of area for some habitats. In addition, habitat modification will not follow a regular pattern over the projected sea levels. Two of the headlands included in the study currently have the maximum level of protection within the SIMP. However, these headlands are likely to lose much of the habitat known to support biodiverse assemblages and may not continue to be suitable sanctuaries into the future. The fine-scale approach taken in this study thus provides a protocol not only for modelling habitat modification but also for future proofing conservation measures under a scenario of changing sea levels.     

Cereal density and N-fertiliser effects on the flora and biodiversity value of arable headlands

Modern intensive farming caused pronounced changes to the European arable flora. Many species adapted to less intensive traditional farming declined severely, as did the potential of unsown arable vegetation to support higher trophic levels. To reverse these trends, various agri-environment measures were introduced. One such measure is to manage cereal headlands as conservation headlands, involving strict restrictions on pesticide and fertiliser use. An additional modification to management which could reduce crop competition and thus deliver benefits to arable plants is cereal sowing at reduced rates. However, little is known about its benefits to rare and declining arable plants, or to species of value to higher trophic levels, and whether it can be implemented without concomitant increase in undesirable weeds. We set up identical two-factorial experiments in winter wheat and spring barley, combining a nitrogen fertiliser versus no fertiliser treatment with cereal sowing at economic rates versus sowing at rates reduced by 75 %, with added sowing of a mixture of rare arable species. Both experiments also included an uncropped but cultivated control equivalent to another agri-environment measure. Our results show that reduced cereal sowing in conservation headlands can benefit rare and declining species, as well as arable plant diversity, without necessarily resulting in a concomitant increase in undesirable weeds. While such benefits tended to be larger in uncropped cultivated controls, conservation headlands have the advantage of not requiring land being taken out of production. Moreover, as shown in this study, their benefits to arable plants can be maximised by reduced sowing.     

Altitudinal and environmental variation in lifespan in the Copper butterfly Lycaena tityrus

1.  Variation in longevity within and between natural populations is widespread, and understanding the relative importance of environmental and genetic factors as well as their interactions in mediating such variation is crucial in longevity research.
2.  In this study lifespan of adult copper butterflies was examined in relation to altitude, temperature (20 and 27 °C), sex and adult feeding.
3.  As expected, longevity increased with decreasing temperature, and sucrose-fed butterflies had longer lifespans compared to water-fed and finally non-fed individuals. The impact of feeding, especially of having access to water or not, was larger at the higher compared to the lower temperature.
4.  Regarding altitudinal patterns, increased lifespan in high-altitude populations was largely restricted to beneficial feeding conditions, while under carbohydrate deprivation low-altitude animals lived longer, suggesting that low-altitude butterflies do better under food stress.
5.  Differences in longevity between sexes were small at 20 °C, while females lived substantially longer than males at the higher temperature. Consequently, females may be less susceptible to high temperature stress than males. Further, males suffered more from food stress than females, suggesting that females are generally more stress resistant than males.
6.  Using a full factorial design, this study demonstrates that variation in longevity is caused by several factors, and additionally by substantial interactive effects. Consequently, patterns of variation in longevity are complex, and one needs to be cautious when neglecting this source of variation, by focussing on individual factors only.     

Butterfly diversity in Mediterranean islands and in Pentadaktylos <Emphasis Type="Italic">Pinus</Emphasis><Emphasis Type="BoldItalic"> </Emphasis><Emphasis Type="Italic">brutia</Emphasis> forests of Cyprus

We analysed the influence of contemporary geography on butterfly diversity for islands in the Mediterranean Basin. We found that island size and distance from the mainland has a significant effect on the number of species. We also used butterflies as an indicator group to identify the importance of forest habitats for biodiversity conservation in the island of Cyprus. To understand the relative importance of local vegetation characteristics of butterflies in the Pentadaktylos mountains transect counts were used to assess the abundance and butterfly diversity in two different forest types. A total of 1,602 butterflies and 23 species were recorded during this research. We observed highly significant effects of forest type on abundance and species richness of butterflies. For example, number of butterflies was significantly higher in old forest than young pine forest. Also, the abundance of endemic butterflies was highest in old forest habitats. Therefore, the survival of the majority of endemic butterflies in Cyprus may depend on conservation of old forests and their understorey plants.     

Physiological tolerance, climate change, and a northward range shift in the spittlebug, Philaenus spumarius

Abstract.  1. Considerable evidence indicates that the Earth's climate has warmed over the past 20 years and most models predict that this process will continue or accelerate. Previous studies involving butterflies and other species have shown that ranges have shifted towards the poles although the mechanisms responsible for these range shifts have not been demonstrated conclusively.
2. Here it is reported that the range of the meadow spittlebug has moved northward along the California coast since 1988.
3. Survival and reproduction of this species in previous laboratory experiments were very sensitive to humidity and temperature. Small deviations from optimal conditions resulted in high mortality.
4. These laboratory results were corroborated by annual field censuses from one location since 1983 in which population densities of spittlebug nymphs were positively correlated with summer humidity and negatively correlated with temperature deviations from the published optimum. Thus it is shown that there are strong links between physiological tolerance and a geographic range shift associated with climatic change.     

Modelling pest population resurgence due to recolonization of fields following an insecticide application

1. To investigate the effect of insect recolonization on the insecticide-induced resurgence of crop pests, a modified Lotka–Volterra predator–prey model was partitioned into two areas, 'sprayed' and 'unsprayed'.
2. The unsprayed area provided a source of insects to recolonize the sprayed area, resulting in a change in pest dynamics in both areas following an insecticide application.
3. Model sensitivity to insecticide selectivity and rates of predation, insect dispersal and pest population increase were examined.
4. Resurgence risk in the sprayed area increased with increasing pest dispersal rate, but decreased with increasing predator dispersal rate.
5. Pest resurgence could also occur in the unsprayed area, especially when prey dispersal rates were low. The extent of resurgence in the unsprayed area could in some circumstances be worse than in the sprayed area itself.
6. The more efficient and longer-lived the predators, the greater the level of pest resurgence in both areas following insecticide use.
7. More selective insecticides, killing the pest, but not the predator, reduced resurgence provided that the average life-span of the predators was reasonably long. Even highly selective insecticides could cause resurgence of the pest in the unsprayed area.
8. When the prey carrying capacity of the unsprayed area was increased relative to that of the sprayed area, resurgent effects in the unsprayed area were reduced, but could still be significant under some circumstances.     

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.     

Butterfly diversity and seasonality of Ta Phin mountain area (N. Vietnam,Lao Cai province)

Human pressures on the environment are changing spatially and temporally, with profound implications for the planet’s biodiversity. Butterflies are important indicators for environmental change and are a suitable group to detect areas of high conservation concern and prioritize conservation efforts. To obtain data to support urgently-needed conservation measures, we surveyed the butterfly fauna in a mountainous region of northern Vietnam, using transect counts over 8 months (121 survey days) from June 2014 to April 2015. In total, we recorded >26,000 butterflies belonging to 231 species, including at least two species new to Vietnam, three red-listed and protected species listed by CITES. Most species were rare: we recorded members of 100 species ≤5 times and 52 species just once. Males dominated the sample, accounting for 81% of all observed butterflies and all members of 84 observed species. Species richness and abundance were highest in July, and there were surprisingly large changes in species composition between months. Species richness curves reached saturation, indicating that we detected most species present in the area, except for members of two families (Lycaenidae and Hesperiidae). Our results highlight the importance of thorough studies using standardized methods, capable of detecting most species in an area, over a whole season. There are urgent needs to integrate butterflies into conservation programs and use their potential as indicator species of habitat degradation and land use intensity.     

Discrimination and release of unpalatable butterflies by Nephila clavipes, a neotropical orb-weaving spider

Abstract. 1. Nephila clavipes (L.), a common spider in neotropical forests, discriminates some unpalatable prey and releases them unharmed from its web. Release is not accidental but results from a specific behavioural sequence.
2. Field trials with twenty-seven butterfly species showed that spiders respond consistently to butterfly species and higher taxa. Ithomiinae and some Danainae are almost always released while Heliconiinae, Nymphalinae, Acraeinae, Pieridae and Papilionidae are usually eaten.
3. Paired tests showed that an immediately preceding experience with a different butterfly did not reveal any change in the spider's usual response to a particular butterfly.
4. Warning coloration is not involved in spider response. Spiders rejected the models but ate the mimics of two different butterfly species pairs. Distastefulness is probably signalled by chemical cues.
5. Some unpalatable butterflies stay motionless when entangled and while the spiders release them. Motionlessness in webs seems to be a requisite to allow recognition of their distastefulness without being bitten by the spider.
6. Spiders differ from vertebrates in the prey accepted and in rejection mechanics. Although there is no indication of learning, spiders may select butterflies for distastefulness, chemical signals and motionlessness, at the individual level.