Using a phenological network to assess weather influences on first appearance of butterflies in the Netherlands

Phenological responses of butterflies to temperature have been demonstrated in several European countries by using data from standardized butterfly monitoring schemes. Recently, phenological networks have enabled volunteers to record phenological observations at project websites. In this study, the quality of the first appearance data of butterflies from the Dutch phenological network ‘De Natuurkalender’ was examined and these data were then used to analyze trends in butterfly appearance between 2001 and 2013, the effects of climatic factors on appearance of butterflies as well as the phenological interaction of one butterfly species, Anthocharis cardamines, and its two major host plants. Although phenological networks are relatively unstructured, this study shows that data from De Natuurkalender were highly comparable to more standardized data collected by the Dutch Butterfly Monitoring Scheme. No trend in first appearance of any butterfly species was found during the time period 2001–2013. The first appearance dates of most butterflies showed, however, a clear relationship with spring temperature. Higher temperatures, especially in March and April, advanced the first appearance of butterflies. Therefore, with climatic warming in the future, earlier appearance of butterflies is expected. Although climate warming is a potential threat for phenological mismatches between different trophic levels, this study shows a similar temperature response of A. cardamines and its main host plants in the Netherlands. However, as only few phenological interactions between species are examined, further research including rarer monophagous butterfly species and their host plants is needed.     

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.     

Altered geographic and temporal variability in phenology in response to climate change

Aim In response to recent climate warming, numerous studies have reported an earlier onset of spring and, to a lesser degree, a later onset of autumn, both determined from phenological observations. Here, we examine whether these reported changes have affected the synchronization of events on a regional level by examining temporal and spatial variability in phenology. In particular, we study whether years with earlier springs are associated with an altered spatial variability in phenology. Location Germany and the United Kingdom. Methods Plant phenological observations of 35 different phases (events such as flowering and leafing) collected by the German Weather Service (1951–2002) and butterfly phenological records of 29 species collected by the UK Butterfly Monitoring Scheme (1976–2003) are used. In these long‐term records, we examine the temporal (year‐to‐year) variability and the spatial (geographic or between site) variability with particular emphasis on how they vary with time of the year and with earliness or lateness of the phase. Results Early phenological events (i.e. spring) are more variable than later events, both in time and in space, although the pattern is clearer for plants than for butterflies. Confirming previous results, we find a clear relationship between the mean date of spring and summer phases and the degree to which they have become earlier. The spatial variability of spring events is greater in warmer years that have faster plant development. However, late spring and summer events do not show a consistent relationship. Autumn events are somewhat more spatially variable in years characterized by later seasons. Main conclusions This is the first examination of spatial variability of plant and animal phenological events at a multinational scale. Earlier spring events are likely to be associated with increased spatial variability in plants, although this is unlikely to also be true for summer events. If species experience differential changes in geographic variation this may disrupt interactions among them, e.g. in food webs. On the other hand, these may offer advantages for mobile species. Further research on linked species is recommended.     

Trait matching and phenological overlap increase the spatio‐temporal stability and functionality of plant–pollinator interactions

Morphology and phenology influence plant–pollinator network structure, but whether they generate more stable pairwise interactions with higher pollination success remains unknown. Here we evaluate the importance of morphological trait matching, phenological overlap and specialisation for the spatio‐temporal stability (measured as variability) of plant–pollinator interactions and for pollination success, while controlling for species' abundance. To this end, we combined a 6‐year plant–pollinator interaction dataset, with information on species traits, phenologies, specialisation, abundance and pollination success, into structural equation models. Interactions among abundant plants and pollinators with well‐matched traits and phenologies formed the stable and functional backbone of the pollination network, whereas poorly matched interactions were variable in time and had lower pollination success. We conclude that phenological overlap could be more useful for predicting changes in species interactions than species abundances, and that non‐random extinction of species with well‐matched traits could decrease the stability of interactions within communities and reduce their functioning.     

Effects of climatic change on the phenology of butterflies in the northwest Mediterranean Basin

Phenological changes in response to climatic warming have been detected across a wide range of organisms. Butterflies stand out as one of the most popular groups of indicators of climatic change, given that, firstly, they are poikilothermic and, secondly, have been the subject of thorough monitoring programmes in several countries for a number of decades. Here we provide for the first time strong evidence of phenological change as a consequence of recent climatic warming in butterflies at a Spanish site in the northwest Mediterranean Basin. By means of the widely used Butterfly Monitoring Scheme methodology, three different phenological parameters were analysed for the most common species to test for trends over time and relationships with temperature and precipitation. Between 1988 and 2002, there was a tendency for earlier first appearance dates in all 17 butterfly species tested, and significant advances in mean flight dates in 8 out of 19 species. On the other hand, the shape of the curve of adult emergence did not show any regular pattern. These changes paralleled an increase of 1–1.5°C in mean February, March and June temperatures. Likewise, a correlation analysis indicated the strong negative effect of spring temperature on phenological parameters (i.e. higher temperatures tended to produce phenological advances), and the opposite effect of precipitation in certain months. In addition, there was some evidence to indicate that phenological responses may differ between taxonomic lineages or species with similar diets. We discuss the consequences that these changes may have on species' population abundances, especially given the expected increase in aridity in the Mediterranean Basin caused by current climatic warming. We predict that varying degrees of phenological flexibility may account for differences in species' responses and, for multivoltine species, predict strong selection favouring local seasonal adaptations such as diapause phenomena or migratory behaviour.     

The conceptual and practical implications of interpreting diet breadth mechanistically in generalist predatory insects

Seasonal polyphenism in animal colour patterns indicates that temporal variation in selection pressures maintains phenotypic plasticity. Spring generation of the polyphenic European map butterfly Araschnia levana has an orange–black fritillary‐like pattern whilst individuals of the summer generation are black with white bands across the wings. What selects for the colour difference is unknown. Because predation is a major selection pressure for insect coloration, we first tested whether map butterfly coloration could have a warning function (i.e. whether the butterflies are unpalatable to birds). In a following field experiment with butterfly dummies we tested whether the spring form is better protected than the summer form from predators in the spring, and vice versa in the summer. The butterflies were palatable to birds (blue tits Cyanistes caeruleus) and in the field the spring and summer form dummies were attacked equally irrespective of season. Therefore, we found no evidence that the map butterfly is warning‐coloured or that seasonal polyphenism is an adaptation to avian predation. Because insect coloration has multiple functions and map butterfly coloration is linked to morphology, life history and development it is likely that the interplay of several selection pressures explains the evolution of colour polyphenism. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Direct and indirect effects of weather variability in a specialist butterfly

1. Climate change poses serious threats to the long‐term persistence of many animal and plant populations. Species having specific niche requirements, or characterised by highly co‐evolved interactions, will face the greatest challenges. An example is represented by Maculinea alcon (Denis & Schiffermüller), a monophagous and univoltine butterfly species, which lays eggs only on larval host plants which occur inparticular phenological conditions. 2. The present 2‐year study focused on two M. alcon populations, both located at the southern boundaries of the species, but facing different climatic conditions (360 m, low altitude versus 860 m, high altitude). Population vulnerability with respect to direct and indirect effects of climate change was analysed, focusing on two important aspects of butterfly biology, i.e. the flight activity of adults and the degree of synchrony in the larval plant–insect interactions. 3. It was observed that, when positive temperature anomalies are reached, the temperature can exert detrimental effects on adults' activity. At a low altitude, in a hotter than usual year, a temperature threshold was recorded (around 32 °C), above which the activity of butterflies is inhibited. In contrast, at a high altitude, temperature increases maintain the opportunity to enhance butterfly activity. Altitudinal differences were also observed in the phenology of the two interacting species, which generate stronger asynchrony at low altitudes. 4. High‐ and low‐altitude populations represent different conservation units: a global increase in temperature would pose a serious threat to the lowland populations, whereas high‐altitude populations would gain a greater role in assuring the persistence of M. alcon at its southern boundaries.     

Differential Bird Predator Attack Rate on Seasonal Forms of the Map Butterfly (Araschnia levana L.): Does the Substrate Matter?

The European map butterfly (Araschnia levana L.) is a striking example of seasonal plasticity. Individuals of the spring generation are reddish with a fritillary‐like colour pattern, whereas the summer generation is black with a white dorsal stripe. Proximate factors explaining the development of the forms are well known, but ultimate explanations have not been tested experimentally yet. The reddish spring form is assumed to have a warning coloration, as found in other nymphalid butterflies that are unpalatable (Aglais urticae). We tested for differential predation by a visually hunting predator (Parus major) in a laboratory experiment using artificial butterflies designed to represent the spring and summer form. Birds were released individually in a flight cage where the alternative forms were presented. Summer forms were more frequently attacked than spring forms, which may point to some aversion against the reddish spring form. But there was also a strong effect of the interaction between seasonal form and type of substrate. Spring forms were much better protected from attacks on the brown substrate of dead leaves compared to the green substrate of nettle leaves. On the latter substrate, latency times before attacking spring forms were on average 2.5 times longer than for summer forms. Experiments with artificial butterflies simplify complex predator–prey interactions because they exclude potential taste or odour effects and they also exclude behavioural responses and interactions of the butterfly. However, our results based on static visual signals provide a promising first step to test the functional significance of this striking seasonal polyphenism.     

Climate changes affecting biotic interactions,phenology, and reproductive success in a savanna community over a 10-year period

Climatic parameters are able to influence the timing of phenological events affecting the degree of synchrony among plant species, their interactions, and reproductive success. Shrubs of Malpighiaceae family in the Brazilian Tropical Savanna present sequential flowering phenology. We verified variations in climatic factors (temperature and precipitation) over a period of 10 years (2005–2014) and correlated them with the onset of flowering of four of these Malpighiaceae species. Furthermore, we tested whether the phenological synchronization among species has changed over time affecting the herbivory and fruit set. Herbivory and fruit production were recorded during three reproductive seasons (2008/2009, 2011/2012, 2013/2014). We developed a mathematical model to estimate the flower and fruit production in response to phenological changes for the next 5 years. Results show that climatic factors changed, influencing the onset of species flowering. The degree of overlap among species also changed and the effects on species interactions were species specific. The mathematical model successfully presented a tendency on flower and fruit production contributing to the predictions of the outcomes in response to phenological changes. We confirm the effects of climate changes on plant phenological events and the importance of feature plasticity for better performance of species.     

Effects of decadal climate change on zooplankton over the last 50 years in the western subarctic North Pacific

Decadal‐ to multi‐decadal variations have been reported in many regional ecosystems in the North Pacific, resulting in an increasing demand to elucidate the link between long‐term climatic forcing and marine ecosystems. We detected phenological and quantitative changes in the copepod community in response to the decadal climatic variation in the western subarctic North Pacific by analyzing the extensive zooplankton collection taken since the 1950s, the Odate Collection. Copepod species were classified into five seasonal groups depending on the timing of the annual peak in abundance. The abundance of the spring community gradually increased for the period 1960–2002. The spring–summer community also showed an increasing trend in May, but a decadal oscillation pattern of quasi‐30‐year cycles in July. Phenological changes coincided with the climate regime shift in the mid‐1970s, indicated by the Pacific decadal oscillation index (PDO). After the regime shift, the timing of the peak abundance was delayed one month, from March–April to April–May, in the spring community, whereas it peaked earlier, from June–July to May–June, in the spring–summer community, resulting in an overlap of the high productivity period for the two communities in May. Wintertime cooling, followed by rapid summertime warming, was considered to be responsible for delayed initiation and early termination of the productive season after the mid‐1970s. Another phenological shift, quite different from the previous decade, was observed in the mid‐1990s, when warm winters followed by cool summers lengthened the productive season. The results suggest that climatic forcing with different decadal cycles may operate independently during winter–spring and spring–summer to create seasonal and interannual variations in hydrographic conditions; thus, combinations of these seasonal processes may determine the annual biological productivity.     

Phenological shifts and the fate of mutualisms

Climate change is altering the timing of life history events in a wide array of species, many of which are involved in mutualistic interactions. Because many mutualisms can form only if partner species are able to locate each other in time, differential phenological shifts are likely to influence their strength, duration and outcome. At the extreme, climate change‐driven shifts in phenology may result in phenological mismatch: the partial or complete loss of temporal overlap of mutualistic species. We have a growing understanding of how, when, and why phenological change can alter one type of mutualism–pollination. However, as we show here, there has been a surprising lack of attention to other types of mutualism. We generate a set of predictions about the characteristics that may predispose mutualisms in general to phenological mismatches. We focus not on the consequences of such mismatches but rather on the likelihood that mismatches will develop. We explore the influence of three key characteristics of mutualism: 1) intimacy, 2) seasonality and duration, and 3) obligacy and specificity. We predict that the following characteristics of mutualism may increase the likelihood of phenological mismatch: 1) a non‐symbiotic life history in which co‐dispersal is absent; 2) brief, seasonal interactions; and 3) facultative, generalized interactions. We then review the limited available data in light of our a priori predictions and point to mutualisms that are more and less likely to be at risk of becoming phenologically mismatched, emphasizing the need for research on mutualisms other than plant–pollinator interactions. Future studies should explicitly focus on mutualism characteristics to determine whether and how changing phenologies will affect mutualistic interactions.     

Lepidoptera rhopalocera in the core of the transboundary biosphere territory “Altai:” Peculiarities of the fauna and population

Migration pathways of butterflies in Southeast Altai (the core of the Transboundary biosphere territory “Altai”) in the spring and summer seasons of 1990 and over the period of 1995–2006 were analyzed. The density of the population was assessed, and specific features of the butterfly distribution among four high-altitude belts (alpine tundra, subalpine, forest, and steppe) were revealed. Eighty-five habitats were examined and butterfly migration pathways were recorded for 359 hours of route surveys with a total length of 875 km. More than 16.5 thousand individuals belonging to 6 families, 66 genera and 107 species were recorded. The total species abundance of butterflies was found to be maximal in the forest belt; the highest population density was recorded in the alpine-tundra belt. The results obtained indicate that the basic differences among the butterfly populations are determined by the thermal conditions related to the altitude and, to a lesser degree, the degree of forestation. The distribution of most butterfly species over the transboundary territory is primarily determined by the recent environmental conditions, in particular the aridity of the tundra-steppe belt.     

The influence of life history traits on the phenological response of British butterflies to climate variability since the late‐19th century

Many species of plants and animals have advanced their phenology in response to climate warming in recent decades. Most of the evidence available for these shifts is based on data from the last few decades, a period coinciding with rapid climate warming. Baseline data is required to put these recent phenological changes in a long‐term context. We analysed the phenological response of 51 resident British butterfly species using data from 83 500 specimens in the collections of the Natural History Museum, London, covering the period 1880–1970. Our analysis shows that only three species significantly advanced their phenology between 1880 and 1970, probably reflecting the relatively small increase in spring temperature over this period. However, the phenology of all but one of the species we analysed showed phenological sensitivity to inter‐annual climate variability and a significant advancement in phenology in years in which spring or summer temperatures were warm and dry. The phenologies of butterfly species were more sensitive to weather if the butterfly species was early flying, southerly distributed, and a generalist in terms of larval diet. This observation is consistent with the hypothesis that species with greater niche breadth may be more phenologically sensitive than species with important niche constraints. Comparison of our results with post‐1976 data from the UK Butterfly Monitoring Scheme show that species flying early in the year had a greater rate of phenological advancement prior to the mid‐1970s. Additionally, prior to the mid‐1970s, phenology was influenced by temperatures in March or April, whereas since 1976, February temperature had a stronger influence on the phenology. These results suggest that early flying species may be approaching the limits of phenological advancement in response to recent climate warming.     

Temporal asynchrony in fine‐root biomass may contribute to shrub and grass coexistence in mixed patches

We described seasonal changes in fine‐root biomass of a grass and a shrub dominant species in a plant community characteristic of the arid Patagonian Monte and then we inferred to want extent the observed differences could contribute to the species coexistence. We selected representative plant patches of the natural vegetation arrangement consisting of one isolated plant of the dominant shrub Larrea divaricata (Ld), grass patches formed by one or more bunches of the dominant grass Nassella tenuis (Nt), and mixed patches consisting of one individual of L. divaricata with bunches of N. tenuis under its canopy (LdNt). We assessed the biomass and temporal changes in fine roots of each species in the upper soil (50 cm depth) of each patch type at three‐month intervals during 2 years. The temporal series of fine‐root biomass were compared among patch types and in relation to above‐ground phenology, as well as climate variables (precipitation, arid index and air temperature). Seasonal changes in fine‐root biomass showed similar cycles in the three plant patches with a maximum in spring. The maximum increase in root biomass in Ld and Nt patches occurred during the onset of reproductive growth in winter and spring, respectively. Fine‐root changes in LdNt patches mimicked that in Ld patches. Precipitation inputs were significantly positively and temperature negatively related to fine‐root changes in Nt patches. Fine‐root changes in Ld and LdNt patches were related to the aridity index (positively) and temperature (negatively). We concluded that the observed asynchronies in the date of the largest increases in root biomass and its climate control between the studied grass and shrub species could contribute to the coexistence of plants of both life forms when they overlap their root systems growing in mixed patches. Mechanisms underlying the root patterns observed should be further explored.     

Climate‐based empirical models show biased predictions of butterfly communities along environmental gradients

A better understanding of the factors that mould ecological community structure is required to accurately predict community composition and to anticipate threats to ecosystems due to global changes. We tested how well stacked climate‐based species distribution models (S‐SDMs) could predict butterfly communities in a mountain region. It has been suggested that climate is the main force driving butterfly distribution and community structure in mountain environments, and that, as a consequence, climate‐based S‐SDMs should yield unbiased predictions. In contrast to this expectation, at lower altitudes, climate‐based S‐SDMs overpredicted butterfly species richness at sites with low plant species richness and underpredicted species richness at sites with high plant species richness. According to two indices of composition accuracy, the Sorensen index and a matching coefficient considering both absences and presences, S‐SDMs were more accurate in plant‐rich grasslands. Butterflies display strong and often specialised trophic interactions with plants. At lower altitudes, where land use is more intense, considering climate alone without accounting for land use influences on grassland plant richness leads to erroneous predictions of butterfly presences and absences. In contrast, at higher altitudes, where climate is the main force filtering communities, there were fewer differences between observed and predicted butterfly richness. At high altitudes, even if stochastic processes decrease the accuracy of predictions of presence, climate‐based S‐SDMs are able to better filter out butterfly species that are unable to cope with severe climatic conditions, providing more accurate predictions of absences. Our results suggest that predictions should account for plants in disturbed habitats at lower altitudes but that stochastic processes and heterogeneity at high altitudes may limit prediction success of climate‐based S‐SDMs.     

Temperature cues phenological synchrony in ant‐mediated seed dispersal

Species‐specific climate responses within ecological communities may disrupt the synchrony of co‐evolved mutualisms that are based on the shared timing of seasonal events, such as seed dispersal by ants (myrmecochory). The spring phenology of plants and ants coincides with marked changes in temperature, light and moisture. We investigate how these environmental drivers influence both seed release by early and late spring woodland herb species, and initiation of spring foraging by seed‐dispersing ants. We pair experimental herbaceous transplants with artificial ant bait stations across north‐ and south‐facing slopes at two contrasting geographic locations. This use of space enables robust identification of plant fruiting and ant foraging cues, and the use of transplants permits us to assess plasticity in plant phenology. We find that warming temperatures act as the primary phenological cue for plant fruiting and ant foraging. Moreover, the plasticity in plant response across locations, despite transplants being from the same source, suggests a high degree of portability in the seed‐dispersing mutualism. However, we also find evidence for potential climate‐driven facilitative failure that may lead to phenological asynchrony. Specifically, at the location where the early flowering species (Hepatica nobilis) is decreasing in abundance and distribution, we find far fewer seed‐dispersing ants foraging during its fruit set than during that of the later flowering Hexastylis arifolia. Notably, the key seed disperser, Aphaenogaster rudis, fails to emerge during early fruit set at this location. At the second location, A. picea forages equally during early and late seed release. These results indicate that climate‐driven changes might shift species‐specific interactions in a plant–ant mutualism resulting in winners and losers within the myrmecochorous plant guild.     

Predicting insect phenology across space and time

Many species appear to be undergoing shifts in phenology, arising from climate change. To predict the direction and magnitude of future changes requires an understanding of how phenology depends on climatic variation. Species show large‐scale spatial variation in phenology (affected by differentiation among populations) as well as variation in phenology from year‐to‐year at the same site (affected predominantly by local plasticity). Teasing apart spatial and temporal variation in phenology should allow improved predictions of phenology under climate change. This study is the first to quantify large‐scale spatial and temporal variation in the entire emergence pattern of species, and to test the relationships found by predicting future data. We use data from up to 33 years of permanent transect records of butterflies in the United Kingdom to fit and test models for 15 butterfly species. We use generalized additive models to model spatial and temporal variation in the distribution of adult butterflies over the season, allowing us to capture changes in the timing of emergence peaks, relative sizes of peaks and/or number of peaks in a single analysis. We develop these models using data for 1973–2000, and then use them to predict phenologies from 2001 to 2006. For six of our study species, a model with only spatial variation in phenology is the best predictor of the future, implying that these species have limited plasticity. For the remaining nine species, the best predictions come from a model with both spatial and temporal variation in phenology; for four of these, growing degree‐days have similar effects over space and time, implying high levels of plasticity. The results show that statistical phenology models can be used to predict phenology shifts in a second time period, suggesting that it should be feasible to project phenologies under climate change scenarios, at least over modest time scales.     

Host plant availability potentially limits butterfly distributions under cold environmental conditions

Species ranges are shaped by both climatic factors and interactions with other species. The stress gradient hypothesis predicts that under physiologically stressful environmental conditions abiotic factors shape range edges while in less stressful environments negative biotic interactions are more important. Butterflies provide a suitable system to test this hypothesis since larvae of most species depend on biotic interactions with a specific set of host plants, which in turn can shape patterns of occurrence and distribution. Here we modelled the distribution of 92 butterfly and 136 host plant species with three different modelling algorithms, using distribution data from the Swiss biodiversity monitoring scheme at a 1 × 1 km spatial resolution. By comparing the ensemble prediction for each butterfly species and the corresponding host plant(s), we assessed potential constraints imposed by host plant availability on distribution of butterflies at their distributional limits along the main environmental gradient, which closely parallels an elevational gradient. Our results indicate that host limitation does not play a role at the lower limit. At the upper limit 50% of butterfly species have a higher elevational limit than their primary host plant, and 33% have upper elevational limits that exceed the limits of both primary and secondary hosts. We conclude that host plant limitation was not relevant to butterfly distributional limits in less stressful environments and that distributions are more likely limited by climate, land use or antagonistic biotic interactions. Obligatory dependency of butterflies on their host plants, however, seems to represent an important limiting factor for the distribution of some species towards the cold, upper end of the environmental gradient, suggesting that biotic factors can shape ranges in stressful environments. Thus, predictions by the stress gradient hypothesis were not always applicable.     

Phenology of British butterflies and climate change

Data from a national butterfly monitoring scheme were analysed to test for relationships between temperature and three phenological measures, duration of flight period and timing of both first and peak appearance. First appearances of most British butterflies has advanced in the last two decades and is strongly related to earlier peak appearance and, for multibrooded species, longer flight period. Mean dates of first and peak appearance are examined in relation to Manley's central England temperatures, using regression techniques. We predict that, in the absence of confounding factors, such as interactions with other organisms and land‐use change, climate warming of the order of 1 °C could advance first and peak appearance of most butterflies by 2–10 days.     

Who flies first? – habitat‐specific phenological shifts of butterflies and orthopterans in the light of climate change: a case study from the south‐east Mediterranean

1. Insects undergo phenological change at different rates, showing no consistent trend between habitats, time periods, species or groups. Understanding how and why this variability occurs is crucial. 2. Phenological patterns of butterflies and Orthoptera were analysed using a novel approach of standardised major axis (SMA) analysis. It was investigated whether: (i) phenology (the mean date and duration of flight) of butterflies and Orthoptera changed from one survey (1998 and 1999 respectively) to another (2011), (ii) the rate at which phenology changed differed between taxa and (iii) phenological change was significantly different across habitat types (agriculture fields, grasslands, and forests). Using the 2011 dataset, we investigated relationships between habitat‐specific variables and species phenology. 3. For both groups, late‐emerging species had an advanced onset on the second survey while the duration showed no consistent trend for butterflies and did not change for Orthoptera. Although the rate at which phenology changed was consistent between the two groups, at the habitat level, a longer duration of flight period emerged for butterflies in agriculture fields while Orthoptera showed no differentiation in flight duration between habitats. We found an earlier emergence of butterflies in grasslands compared to forests, attributed to habitat‐specific temperature, whereas spatial variation in humidity had a significantly lower effect on butterflies' phenology in grasslands compared to forests. A gradual delay of butterfly appearances as the canopy cover increased was also found. 4. The utility of SMA analysis was demonstrated in phenological studies and evidence was detected that both habitat type and habitat‐specific variables refine species' phenological responses.