Geographical variability in, and temperature effects on, the phenology of Maniola jurtina and Pyronia tithonus (Lepidoptera, Satyrinae) in England and Wales

ABSTRACT. 1. Geographical variability in, and temperature effects on, the mean date of adult flight period and the SD about this date are analysed for two univoltine, grassland butterflies in England and Wales from 1976 to 1985. Data were collected on the Butterfly Monitoring Scheme for Maniola jurtina (L.) at twenty-nine sites and Pyronia tithonus (L.) at twenty sites.
2. Substantial variability for mean date and SD occurs between years and between sites. Changes in mean date between years tend to occur consistently at different sites. The species show some parallel in variation between sites, especially for mean date.
3. June maximum temperature accounts for 95% and 75% of the variation in mean date between years in M. jurtina and P. tithonus , respectively ( r =-0.97 and -0.87). Similar relationships occur for temperatures cumulated over the period of post-winter development from March to July or August.
4. Greater geographical variability in phenology, and a generally less synchronized flight period in M.jurtina may be associated with broader habitat preferences than in P. tithonus.
5. The mean date of adult flight period remains at roughly the same date at more northerly latitudes. In M.jurtina the flight period becomes more synchronized, begins later and ends earlier in the north. P. tithonus shows little or no indication of such a response to latitude. This is discussed with regard to changes in season length and factors limiting the species'range.     

Comparative phenology of four satyrine butterflies inhabiting dry grasslands in Sweden

The phenology of the satyrine butterflies Coenonympha pamphilus L., Lasiommata megera L., Hipparchia semele L. and Maniola jurtina L. was compared under seminatural conditions with Festuca ovina as the food-plant. Coenonympha pamphilus shows striking phenotypic plasticity as revealed by initiating cohorts at successive dates during the whole flight period. Growth rate is finely tuned depending on season, and larvae can hibernate in any of the three first instars, but also aestivate when 'surplus'time is available. Hipparchia semele and M. jurtina have reduced growth rates as last instar larvae at the beginning of summer. Lasiommata megera , which has an obligate bivoltine life cycle, shows the highest overall growth rate of all four species. Results suggest that egg size variation has consequences for the life cycles of these species.     

Escape from natural enemies during climate-driven range expansion: a case study

Abstract.  1. A major, and largely unexplored, uncertainty in projecting the impact of climate change on biodiversity is the consequence of altered interspecific interactions, for example between parasitoids and their hosts. The present study investigated parasitism in the Brown Argus butterfly, Aricia agestis ; a species that has expanded northward in Britain during the last 30  years in association with climate warming.
2.  Aricia agestis larvae suffered lower mortality from parasitoids in newly colonised areas compared with long-established populations. This result was consistent over four consecutive generations (2 years) when comparing one population of each type, and also when several populations within the historical and recently colonised range of the species were compared within a single year. Thus, A. agestis appears to be partially escaping from parasitism as it expands northwards.
3. Reduced parasitism occurred despite the fact that several of the parasitoid species associated with A. agestis were already present in the newly colonised areas, supported predominantly by an alternative host species, the Common Blue butterfly, Polyommatus icarus .
4. As the species expand their distributions into areas of increased climatic suitability, invasion fronts may escape from natural enemies, enhancing rates of range expansion. The results suggest that the decoupling of interspecific interactions may allow some species to exploit a wider range of environments and to do so more rapidly than previously thought possible.     

Asymmetric dispersal and survival indicate population sources for grassland butterflies in agricultural landscapes

We tested the hypothesis that populations in small habitat fragments remaining in agricultural landscapes are maintained by repeated immigration, using three grassland butterflies ( Aphantopus hyperantus , Coenonympha pamphilus and Maniola jurtina ). Transect counts in 12 matched sets of semi-natural pastures, and linear habitat elements proximate and isolated from the pastures showed that population densities of M.  jurtina and C.  pamphilus were significantly higher in pastures and in linear habitats adjacent to these than in isolated linear elements. A mark-recapture study in a 2×2 km landscape indicated that individuals of all three species are able to reach even the isolated linear elements situated at least 1 km from the grasslands. For two of the species, A.  hyperantus and C.  pamphilus , analysis of the mark-recapture data revealed higher daily local survival rates in the semi-natural pastures and more individuals dispersing from pastures to linear habitat elements. The proportion of old compared to young individuals of C.  pamphilus and M.  jurtina were significantly higher in linear elements than in semi-natural pastures, which suggests that butterflies emerging in pastures subsequently dispersed to the linear elements. In combination, these results suggest that semi-natural pastures act as population sources, from which adult butterflies disperse to surrounding linear elements. Hence, preservation of the remaining fragments of semi-natural grassland is necessary to keep the present butterfly abundance in the surrounding agricultural landscape.     

Dynamics in a butterfly–plant–ant system: influence of habitat characteristics on turnover rates of the endangered lycaenid Maculinea alcon

Abstract.  1.  Maculinea alcon , a myrmecophilic, stenotopic lycaenid butterfly is restricted to wet heathlands, bogs, and nutrient-poor hay meadows. Due to intensification of agriculture and decrease of extensively grazed meadows, many suitable habitats have disappeared and the remaining ones are highly fragmented and deteriorated.
2. Historical distribution data and a comparison with the present occupation of patches show the decline of this critically endangered butterfly in north-west Germany. Most of the populations in north-west Germany are small and often geographically isolated.
3. In summer 2002, two-thirds of 77 investigated potential patches were unoccupied as a result of unsuitable habitat structure and habitat fragmentation.
4. Several habitat parameters were highly significantly correlated with the presence of M. alcon , in particular the distribution pattern of the host plant Gentiana pneumonanthe. Furthermore, butterflies were absent from many patches with an incidence probability below 50% with respect to patch size and isolation.
5. In the nature reserve Lüneburger Heide, part of the study area, M. alcon populations have been observed since 1995. Typical turnover of local populations could be detected during these years. Extinctions and re-colonisations have stabilised the presence of this species in a metapopulation in this nature reserve.
6. These data show the importance of different factors on different spatial levels influencing the presence of this endangered butterfly.     

Movement of two grassland butterflies in the same habitat network: the role of adult resources and size of the study area

Abstract. 1.  Movement patterns of two butterfly species (meadow brown Maniola jurtina L. and scarce copper Lycaenae virgaureae L.) were studied in a 172 ha area within a landscape with a high percentage of suitable habitats for mark–release–recapture experiments.
2.  Adult resource density, but not patch size or larval food plant abundance, influenced the numbers and the fractions of residents, emigrants, and immigrants.
3.  Differences between species were observed in movement frequency and maximum distances moved but not in mean distances moved.
4.  The scarce copper showed much greater movement ability than expected from the results of published studies. This is believed to be a result of the comparatively large size of the study area and the high cover of suitable habitat (>50%).
5.  The mean and maximum distances travelled by butterflies reflected differences in the size of the study area.     

Cost of reproduction in males of a satyrine butterfly Lethe diana

Adult butterflies feed largely on floral nectar and tree sap, both of which consist mainly of carbohydrates and include little nitrogen. They depend on the larval diet for nitrogenous resources. Consequently, there is a trade‐off between the reproductive and somatic nitrogenous investments of adults. Furthermore, male butterflies invest a considerable amount into spermatophores, containing nitrogen, which they give to their sexual partners. One way in which male butterflies could potentially replenish their spermatophores is by flight muscle histolysis, which may reduce locomotor ability and lifespan. In the present study, the effect of mating experience on nitrogen dynamics and the lifespan of males is investigated in the satyrine butterfly Lethe diana (Butler). Mated males do not have less thoracic nitrogen than virgin males, suggesting that mating experiences do not induce spermatophore recovery through flight muscle histolysis. Mated males possess less abdominal nitrogen than virgin males at death, indicating that they cannot recover the lost nitrogenous resource used for a single mating. Lifespan does not differ between mated and virgin males. Thus, reproduction and longevity are not fuelled necessarily by the same shared resource pools. Spermatophore mass increases as males get older. Considering that resources for producing spermatophores are limited, males may adjust the spermatophore mass strategically, depending on their chances of future reproduction.     

Rewiring of interactions in a changing environment: nettle-feeding butterflies and their parasitoids

Climate and land use change can alter the incidence and strength of biotic interactions, with important effects on the distribution, abundance and function of species. To assess the importance of these effects and their dynamics, studies quantifying how biotic interactions change in space and time are needed. We studied interactions between nettle-feeding butterflies and their shared natural enemies (parasitoids) locally and across 500 km latitudinal gradient in Sweden. We also examined the potential impact of the range-expansion of the butterfly Araschnia levana on resident butterflies via shared parasitoids, by studying how parasitism in resident butterflies covaries with the presence or absence of the newly-established species. We collected 6777 larvae of four nettle-feeding butterfly species (Aglais urticae, Aglais io, Ar. levana and Vanessa atalanta), over two years, at 19 sites distributed along the gradient. We documented the parasitoid complex for each butterfly species and measured their overlap, and analysed how parasitism rates were affected by butterfly species assemblage, variations in abundance, time, and the arrival of Ar. levana. Parasitoids caused high mortality, with substantial overlap in the complex of parasitoids associated with the four host butterflies. Levels of parasitism differed significantly among butterflies and were influenced by the local butterfly species assemblage. Our results also suggest that parasitism in resident butterflies is elevated at sites where Ar. levana has been established for a longer period. In our study system, variations in butterfly species assemblages were associated in a predictable way with substantial variations in rates of parasitism. This relationship is likely to affect the dynamics of the butterfly host species, and potentially cascade to the larger number of species with which they interact. These results highlight the importance of indirect interactions and their potential to reorganise ecological communities, especially in the context of shifts in species distributions in a warmer world.     

Consequences of the arms race between <Emphasis Type="Italic">Maculinea teleius</Emphasis> social parasite and <Emphasis Type="Italic">Myrmica</Emphasis> host ants for myrmecophilous butterfly conservation

The arms race between Maculinea butterflies and Myrmica host ants leads to local host-parasite adaptations. In our study, we assessed whether sympatric and allopatric Myrmica scabrinodis populations exhibit behavioural differences towards Maculinea teleius larvae during the adoption-period when butterfly larvae need to be taken inside the Myrmica nest. The second aim was to assess the butterfly survival rate inside ant colonies from different populations. We used one sympatric host population and three allopatric populations: one infested by M. teleius and two uninfested populations. We found that ants from the sympatric population showed a higher number of positive behaviours toward M. teleius larvae during adoption than ants from the allopatric populations. There were no differences in the number of inspection or negative behaviour events. The survival of butterfly larvae was highest inside sympatric host colonies and differed from the survival of M. teleius reared by ants from the allopatric, uninfested populations. No difference was found for the survival rate of M. teleius raised by infested, allopatric host colonies compared to sympatric host populations. Our results suggest the lack of behavioural counter-adaptations of local hosts of M. teleius that more easily adopt and rear butterfly caterpillars compared to naive M. scabrinodis colonies. Our results may also have implications for Maculinea butterfly conservation, especially for reintroduction programmes. We suggest that the existence of behavioural host defences should be checked for the source host population, as well as for the Myrmica population from the reintroduction site. It may also be reasonable to introduce several Myrmica host colonies from the source butterfly host population.     

Contrasting movement patterns in two species of chequerspot butterflies, Euphydryas aurinia and Melitaea phoebe, in the same patch network

Abstract.  1. Mark–release–recapture studies were conducted on two species of chequerspot butterfly, Euphydryas aurinia and Melitaea phoebe , in the same habitat patch network in Yanjiaping, a small basin in the Taihang Mountains, north-west of Beijing, China, in 2000.
2. Euphydryas aurinia tended to stay in the habitat patches and to move to neighbouring patches, whereas M. phoebe moved widely among patches in the entire network.
3. The parameters of the virtual migration model showed higher daily emigration propensity in M. phoebe and in E. aurinia males than in E. aurinia females, and significantly greater average daily movement distance in M. phoebe than in E. aurinia .
4. The results are consistent with the previous findings showing genetic structuring among local populations of E. aurinia but not among local populations of M. phoebe .
5. Based on the genetic and ecological results, it was concluded that E. aurinia has a classic metapopulation in the study area, whereas M. phoebe appears to have a source–sink metapopulation.
6. In 2000, when there was an overall increase in the abundance of the two species, the limited mobility of E. aurinia resulted in an increase in the average local population size, whereas the increase in the number of local populations in M. phoebe was due to its high mobility .     

Patterns of infestation by the trombiculid mite Eutrombicula alfreddugesi in four sympatric lizard species (genus Tropidurus) in northeastern Brazil

We studied the parasitism by the chigger mite Eutrombicula alfreddugesi on four sympatric lizard species of the genus Tropidurus in Morro do Chapéu, Bahia state, Brazil: T. hispidus, T. cocorobensis, T. semitaeniatus and T. erythrocephalus. For each species, we investigated the patterns of infestation and analyzed to which extent they varied among the hosts. We calculated the spatial niche breadth of the chigger mite on the body of each host species and the distribution of mites along the hosts' bodies for each Tropidurus species. All four species of Tropidurus at Morro do Chapéu were parasited by the chigger mite, with high (97-100%) prevalences. Host body size significantly explained the intensity of mite infestation for all species, except T. erythrocephalus. The body regions with highest intensity of infestation in the four lizard species were the mite pockets. The spacial niche width of the chigger varied consistently among the four lizards species studied being highest for T. erytrocephalus and lowest for T. cocorobensis. We conclude that the distribution and intensity with which lizards of the genus Tropidurus are infested by Eutrombicula alfreddugesi larvae results from the interaction between aspects of host morphology (such as body size and the occurrence and distribution of mite pockets) and ecology (especially microhabitat use).     

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.     

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.     

栖息地质量对两种网蛱蝶集合种群结构和分布的影响

在河北省赤城县研究了栖息地质量对大网蛱蝶Melitaea phoebe和金堇蛱蝶Euphydryas aurinia两种网蛱蝶集合种群结构和分布的影响。这两种网蛱蝶在约10 km_²的区域内共存,成虫期的蜜源植物几乎相同,大网蛱蝶的发生峰期比金堇蛱蝶晚约一个月,两者只有不到一周左右的时间重叠。大网蛱蝶和金堇蛱蝶幼虫的寄主植物分别是: 祁州漏芦（菊科）和华北蓝盆花（川续断科）。蜜源植物的丰度与两种网蛱蝶的局域种群大小呈正相关;祁州漏芦的密度对大网蛱蝶的局域种群大小影响很大,金堇蛱蝶的局域种群大小则与其寄主植物华北蓝盆花的高度正相关;斑块内平均植被高度与两种网蛱蝶的局域种群大小均呈正相关,植物多样性、植物均匀性和植被盖度均与金堇蛱蝶的局域种群大小负相关,与大网蛱蝶的关系不大。同时分析了其他因子如斑块的坡向、坡度等的影响。主要结论是：1）幼虫寄主植物的不同和成蝶飞行峰期的分离允许两种网蛱蝶在这样一个小的斑块区域内共存;2）蜜源是重要的限制因子,并且受气候随机性的影响很大,蜜源的波动可以很好地解释网蛱蝶集合种群在年度间的动态变化;3）大网蛱蝶和金堇蛱蝶的飞行、食物搜寻能力的不同以及各自寄主植物的生物学特性、空间分布的不同决定了它们具有不同的集合种群结构： 金堇蛱蝶是经典的集合种群,大网蛱蝶是源-汇集合种群;4）斑块质量和昆虫行为共同决定了两种网蛱蝶的集合种群结构和分布。     

Poly modal emergence of the tiger swallowtail, Papilio glaucus (Lepidoptera: Papilionidae): source of a false second generation in central New York State

Abstract. 1. The tiger swallowtail butterfly, Papilio glaucus (L.) (Lepidoptera: Papilionidae), is commonly assumed to have a bivoltine life cycle in central New York State and other parts of the northeastern United States. The seasonal abundance of adults in Tompkins County, New York, shows a bimodal pattern in some years, and a skewed unimodal pattern in others. We studied the incidence of diapause, the potential for successful development of two generations in the field, and the dates of adult emergence from overwintering pupae to determine whether these seasonal patterns are the result of bivoltinism.
2. Insects from Tompkins County, reared as larvae under 16L:8D photoperiod, uniformly enter pupal diapause.
3. Comparison of the heat units required for successful development of two generations with the heat units available in the field during 11 years shows that emergence of non-diapausing individuals would occur too late in the summer to contribute to the second peak of butterfly abundance. Offspring of non-diapausing individuals could only rarely complete development before the end of the growth season.
4. Diapausing offspring of butterflies collected in early June and late July differed sufficiently in dates of emergence to account for the field pattern of seasonal abundance.
5. Electrophoretic studies revealed no evidence of genetic isolation between early and late emerging butterflies.
6. These results suggest that polymodal emergence of diapausing insects can masquerade as a bivoltine life cycle.     

Habitat use of the endangered butterfly Euphydryas maturna and forestry in Central Europe

The knowledge of ecological requirements of declining butterflies of European woodlands remains limited, which hinders conservation management of their localities. This also applies for continentally threatened scarce fritillary Euphydryas maturna . On the basis of the largest data set on its habitat use ever collected in Central Europe, we analyse habitat requirements of its populations in Austria (A), the Czech Republic (Cz) and Germany (D). All studied populations inhabit open-canopy sites within woodlands, but larval survival decreases under full sun and preferred sites are relatively humid and sheltered. Nests of pre-hibernation larvae occur at terminal branches of Fraxinus excelsior , 1.5–3 m above the ground. Pre-hibernation mortality reaches 70% (Cz, D). Another limiting factor is quality of woodland vegetation: post-hibernation larvae consume a wide range of herbs and shrubs, and adult distribution is linked to nectar availability. The butterfly thus depends on highly heterogeneous early successional stages of deciduous woods, historically maintained by coppicing (Cz, D) and forest pasture (A). Restoration of these traditional methods offers the only chance for survival of E. maturna in Central Europe, and the butterfly may become a flagship for other threatened organisms of open-canopy woodlands.     

Parasitism of Hylodes phyllodes (Anura: Cycloramphidae) by Hannemania sp (Acari: Trombicuudae) in an area of Atlantic Forest, Ilha Grande, southeastern Brazil

We studied some parameters of the parasitism by the mite Hannemonia sp. on the endemic frog Hylodes phyllodes in the Atlantic Forest of Ilha Grande (Rio de Janeiro State, Brazil). Prevalence, mean abundance, mean intensity and total intensity of infestation, body regions infected, and host sexual differences in parasitism rate of the larvae of Hannemania sp. on individuals of H. phyllodes were assessed. Prevalence was 86.5% (N = 37; total of 1,745 larvae of Hannemania sp) for male hosts and 91.7% (N = 12; total of 330 larvae) for female hosts, with no significant difference between the sexes. Overall prevalence of Hannemania sp. on H. phyllodes was 87.7%. Mean intensity of infestation in males (54.5 +/- 42.5; range 1-173 larvae) was higher than in females (29.9 +/- 47.6; range 1-166). We conclude that the rates of intensity, abundance, and prevalence of Hannemania larvae parasitizing Hylodes phyllodes at Ilha Grande were considerably high, suggesting that this species of anuran constitutes a relevant host for this mite species to complete its life-cycle in the area. Differences between males and females in infestation rates probably reflect their differential use of space in the forest.     

The adaptiveness of searching and host selection behaviour in Pieris rapae (L.)

This study evaluates the adaptive significance of host preferences and searching behaviour in Vancouver and Canberra populations of the cabbage butterfly Pieris rapae (L.). As a result of a complex of responses to plant age, the butterflies concentrate their eggs on middle-aged plants. Young larvae develop faster and survive better on young plants than old ones, but larvae on smaller plants are more susceptible to crowding effects. Thus a preference for plants which are well-grown but not too old is selectively advantageous. By contrast, the butterflies’ host species preferences appear non-adaptive, and are unrelated to the quality of the host as larval food. Vancouver butterflies change their flight direction often and are very responsive to hosts, thereby generating a very aggregated distribution at a low cost in flight time. Canberra butterflies tend to fly in straight lines and are less responsive to hosts; their egg distribution is consequently more nearly random, but they fly further for each egg they lay. The relative costs of aggregation and increased flight time differ between the populations in a manner consistent with the observed behavioural differences.     

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.     

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.