Extended season for northern butterflies

Butterflies are like all insects in that they are temperature sensitive and a changing climate with higher temperatures might effect their phenology. Several studies have found support for earlier flight dates among the investigated species. A comparative study with data from a citizen science project, including 66 species of butterflies in Sweden, was undertaken, and the result confirms that most butterfly species now fly earlier during the season. This is especially evident for butterflies overwintering as adults or as pupae. However, the advancement in phenology is correlated with flight date, and some late season species show no advancement or have even postponed their flight dates and are now flying later in the season. The results also showed that latitude had a strong effect on the adult flight date, and most of the investigated species showed significantly later flights towards the north. Only some late flying species showed an opposite trend, flying earlier in the north. A majority of the investigated species in this study showed a general response to temperature and advanced their flight dates with warmer temperatures (on average they advanced their flight dates by 3.8 days/°C), although not all species showed this response. In essence, a climate with earlier springs and longer growing seasons seems not to change the appearance patterns in a one-way direction. We now see butterflies on the wings both earlier and later in the season and some consequences of these patterns are discussed. So far, studies have concentrated mostly on early season butterfly–plant interactions but also late season studies are needed for a better understanding of long-term population consequences.     

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.     

The effect of spring temperature on the appearance dates of British butterflies 1883–1993

This paper examines the first appearance of butterflies using two sources, historical data from the former phenologieal reports of the Royal Meteorological Society and recent data from the extant Butterfly Monitoring Scheme Using regression techniques mean dates of appearance are related to Manley's central England temperatures and are then examined for other, unexplained, trends over time The observed relationships suggest that, in the absence of evolutionary change, climate warming of the order of 3°C could advance butterfly appearance by two to three weeks The flowering of two of the larval foodplants of the orange tip butterfly is also examined, with the conclusion that synchrony is likely to be maintained by a similar advance in timing The consequences of such changes to the phenology of British butterflies are discussed     

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.     

Natural history museum collections provide information on phenological change in British butterflies since the late-nineteenth century

Museum collections have the potential to provide valuable information on the phenological response of organisms to climate change. This is particularly useful for those species for which few data otherwise exist, but also to extend time series to the period before other observational data are available. To test this potential, we analysed data from 2,630 specimens of four species of British butterflies (Anthocharis cardamines, Hamearis lucina, Polyommatus bellargus and Pyrgus malvae), collected from 1876 to 1999 and stored in the Natural History Museum, London, UK (NHM). In A. cardamines, first-generation P. bellargus and P. malvae, we found that there was a strong significant negative relationship between spring temperature and 10th percentile collection dates, which approximates mean first appearance date, and median collection date, which approximates mean flight date. In all four species, there was a significant negative relationship between the 10th percentile collection date and the length of the collection period, which approximates flight period. In second-generation P. bellargus, these phenological measurements were correlated with summer temperature. We found that the rates of phenological response to temperature, based on NHM data, were similar to, or somewhat greater than, those reported for other organisms based on observational data covering the last 40 years. The lower rate of phenological response, and the significant influence of February rather than March or April temperatures, in recent decades compared with data from earlier in the twentieth century may indicate that early emerging British butterfly species are currently approaching the limits of phenological advancement in response to recent climate warming.     

是否低估了本地热带蝴蝶区系成分的丰富度?样带观察结果的剖析

作者利用在菲律宾吕宋岛Calumpang Lejos (Indang, Cavite) 地区以样带法收集的蝴蝶种数记录, 检验了类群记录是否低估了当地种数丰富度和多样性。在两年间长达340 h的457个样带调查中, 没有出现种数积累曲线突破情况。折叠估计、有偏纪录、丰富度的地区变化以及与群落生境有关的相对多度均支持当地蝴蝶区系丰富度被低估的观点。在样带调查中, 多样性及体色类型记录具有误差。这些误差反映在发现日期以及同种的科学命名日期, 并且涉及到菲律宾吕宋岛发现的所有物种, 且与蝴蝶的体型大小有关。数量较少、体型较小和不显眼的种类发现得较晚。在分类上亦有误差, 弄蝶科和灰蝶科比粉蝶科和蝶科稀少、体小且不显眼, 发现和命名较晚。翅型特征不明显的种类具有体型较小的趋势, 使所做的形态特征记录产生错误。我们发现菲律宾吕宋岛Calumpang Lejos (Indang, Cavite) 地区的稀有种类体型明显较小和特征较不明显。由于区域性样带调查偏差包括了灰蝶科但未包括弄蝶科, 因此偏差源于对树冠型蝴蝶种类的记录偏低。本文讨论了在其它生物区系以及热带蝴蝶保育上的应用, 认为今后需要使用误差小的新技术进行动物区系中物种丰富度的可靠估计。     

Indian migrant butterflies displaced to Arabia by monsoon storm 'Aurora' in August 1983

Although many insect species are now thought to travel hundreds and even thousands of kilometres on the wind (Pedgley, 1982) only a few are butterflies. Three species for which there is reasonably convincing but circumstantial evidence are Danaus plexippus L. from the U.S.A. to the British Isles (Hurst, 1969), Hypolimnas bolina nerina (F.) from Australia to New Zealand (Tomlinson, 1973) and Nymphalis antiopa L. from northern Europe to the British Isles (Chalmers-Hunt, 1977). In this paper we describe the sudden appearance of Indian butterflies in Arabia and provide evidence that they were windborne across the Arabian Sea.     

A reversal of the shift towards earlier spring phenology in several Mediterranean reptiles and amphibians during the 1998–2013 warming slowdown

Herps, especially amphibians, are particularly susceptible to climate change, as temperature tightly controls many parameters of their biological cycle—above all, their phenology. The timing of herps’ activity or migration period—in particular the dates of their first appearance in spring and first breeding—and the shift to earlier dates in response to warming since the last quarter of the 20^th century has often been described up to now as a nearly monotonic trend towards earlier phenological events. In this study, we used citizen science data opportunistically collected on reptiles and amphibians in the northern Mediterranean basin over a period of 32 years to explore temporal variations in herp phenology. For 17 common species, we measured shifts in the date of the species’ first spring appearance—which may be the result of current changes in climate—and regressed the first appearance date against temperatures and precipitations. Our results confirmed the expected overall trend towards earlier first spring appearances from 1983 to 1997, and show that the first appearance date of both reptiles and amphibians fits well with the temperature in late winter. However, the trend towards earlier dates was stopped or even reversed in most species between 1998 and 2013. We interpret this reversal as a response to cooling related to the North Atlantic Oscillation (NAO) in the late winter and early spring. During the positive NAO episodes, for certain species only (mainly amphibians), the effect of a warm weather, which tends to advance the phenology, seems to be counterbalanced by the adverse effects of the relative dryness.     

Vertical stratification of ithomiine butterfly (Nymphalidae: Ithomiinae) mimicry complexes: the relationship between adult flight height and larval host-plant height

Data are presented which confirm previous findings that sympatric mimicry complexes dominated by unpalatable Neotropical ithomiine butterflies (Nymphalidae: Ithomiinae) are vertically stratified by height of flight. Flight height of ithomiine species is positively correlated with the height of their larval host-plants. Thus members of a mimicry complex utilize host-plants of similar heights. Non-mimetic British woodland butterflies also show a positive relationship between flight height and host-plant height, which suggests that the relationship is independent of mimicry. I propose that female butterflies fly at heights which maximize the probability of encountering their larval host-plants, and that males fly at similar heights to females in order to maximize the probability of encountering potential mates. Female butterflies probably encounter plants of similar heights to their larval hosts more frequently than they encounter plants of other heights. I suggest that butterfly species may therefore be more likely to make host shifts to plant species of a similar height to their current host-plants. Finally, I discuss how the relationship between flight height and height of larval host-plants, coupled with microhabitat-dependent selection on colour pattern, could lead to the evolution in sympatry of vertically stratified mimicry complexes.     

Small changes in timing of breeding among subarctic passerines over a 32‐year period

Many bird populations in temperate regions have advanced their timing of breeding in response to a warming climate in recent decades. However, long‐term trends in temperature differ geographically and between seasons, and so do responses of local breeding populations. Data on breeding bird phenology from subarctic and arctic passerine populations are scarce, and relatively little data has been recorded in open‐nesting species. We investigated the timing of breeding and its relationship to spring temperature of 14 mainly open‐nesting passerine species in subarctic Swedish Lapland over a period of 32 years (1984–2015). We estimated timing of breeding from the progress of post‐juvenile moult in mist‐netted birds, a new method exploring the fact that the progress of post‐juvenile moult correlates with age. Although there was a numerical tendency for earlier breeding in most species (on average ?0.09 days/year), changes were statistically significant in only three species (by ?0.16 to ?0.23 days/year). These figures are relatively low compared with what has been found in other long‐term studies but are similar to a few other studies in subarctic areas. Generally, annual hatching dates were negatively correlated with mean temperature in May. This correlation was stronger in long‐distance than in short‐distance migrants. Although annual temperatures at high northern latitudes have increased over recent decades, there was no long‐term increase in mean temperature in May over the study period at this subarctic site. This is probably the main reason why there were only small long‐term changes in hatching dates.     

Temperature‐related shifts in butterfly phenology depend on the habitat

Many species are becoming active earlier in the season as the climate becomes warmer. In parallel to phenological responses to climate change, many species have also been affected by habitat changes due to anthropogenic land use. As habitat type can directly affect microclimatic conditions, concurrent changes in climate and habitat could have interacting effects on the phenology of species. Temperature‐related shifts in phenology, however, have mostly been studied independent of habitat types. Here, I used long‐term data from a highly standardized monitoring program with 519 transects to study how phenology of butterflies is affected by ambient temperature and habitat type. I compared forests, agricultural areas and settlements, reflecting three major land use forms, and considered butterfly species that were observed in all three of these habitats. Seasonal appearance of the butterflies was affected both by the ambient temperature and the habitat type. As expected, warmer temperatures led to an overall advancement of the appearance and flight period of most species. Surprisingly, however, phenology of species was delayed in settlement habitats, even though this habitat type is generally associated with higher temperatures. A possible explanation is dispersal among habitat types, such that source–sink effects affect local phenology. When there is little productivity in settlement areas, observed butterflies may have immigrated from forest or agricultural habitats and thus appear later in settlements. My findings suggest that a spillover of individuals among habitats may affect phenology trends and indicate that phenological studies need to be interpreted in the context of habitat types. This becomes especially important when defining strategies to prevent or mitigate effects of climate and land‐use changes on phenology and abundance of species.     

Spatial and habitat variation in aphid,butterfly, moth and bird phenologies over the last half century

Global warming has advanced the timing of biological events, potentially leading to disruption across trophic levels. The potential importance of phenological change as a driver of population trends has been suggested. To fully understand the possible impacts, there is a need to quantify the scale of these changes spatially and according to habitat type. We studied the relationship between phenological trends, space and habitat type between 1965 and 2012 using an extensive UK dataset comprising 269 aphid, bird, butterfly and moth species. We modelled phenologies using generalized additive mixed models that included covariates for geographical (latitude, longitude, altitude), temporal (year, season) and habitat terms (woodland, scrub, grassland). Model selection showed that a baseline model with geographical and temporal components explained the variation in phenologies better than either a model in which space and time interacted or a habitat model without spatial terms. This baseline model showed strongly that phenologies shifted progressively earlier over time, that increasing altitude produced later phenologies and that a strong spatial component determined phenological timings, particularly latitude. The seasonal timing of a phenological event, in terms of whether it fell in the first or second half of the year, did not result in substantially different trends for butterflies. For moths, early season phenologies advanced more rapidly than those recorded later. Whilst temporal trends across all habitats resulted in earlier phenologies over time, agricultural habitats produced significantly later phenologies than most other habitats studied, probably because of nonclimatic drivers. A model with a significant habitat‐time interaction was the best‐fitting model for birds, moths and butterflies, emphasizing that the rates of phenological advance also differ among habitats for these groups. Our results suggest the presence of strong spatial gradients in mean seasonal timing and nonlinear trends towards earlier seasonal timing that varies in form and rate among habitat types.     

Climatic change is advancing the phenology of moth species in Ireland

Recent increases in global temperatures have contributed to advancing phenology of plants and animals. These increases in temperature have been shown to affect the phenological phases (phenophases) of plants and birds in Ireland, but less is known about the effect on the phenophases of Irish insects. Records of the flight periods of 59 species of Irish moths over the past 35 years (1974–2009) were obtained from a public monitoring group. Observations were analysed across the country using generalized additive models (GAMs) weighted by total yearly population numbers for each species. The results of the statistical analyses showed that 45 of the 59 species studied have a significantly earlier first sighting date now than when observations began. With this earlier emergence, 44 of the 59 species also have a significantly longer flight season over the same 35‐year period. The extent of these changes varies across the country and by species life history. In particular, species emerging in spring are advancing at a much faster rate than species emerging during the summer. Many of these changes in first sighting are negatively correlated with rising temperatures in Ireland, particularly in late spring and early summer (May and June). The variation in phenological advancement in the moth species of Ireland is extremely complex and may be influenced more by species life history than by the phenology of interacting species, such as host plants.     

Museum egg collections as stores of long-term phenological data

Museum collections hold large amounts of data on collecting dates and localities of eggs collected over the past 150 years. Egg collections hold the longest available time series for a wide range of bird species on a large spatial scale. Using data for two British species I investigate whether egg collection data can be used in phenological research. A method is presented allowing laying dates to be estimated from collecting dates. Problems and biases in the data are highlighted. Both the dipper and song thrush have started laying earlier over the past 150 years. The advance in laying is significantly correlated with mean March temperature. Received: 25 October 2000 / Revised: 14 May 2001 / Accepted: 16 May 2001     

Phenological delay despite warming in wood frog Rana sylvatica reproductive timing: a 20-year study

Across all taxa, amphibians exhibit some of the strongest phenological shifts in response to climate change. As climates warm, amphibians and other animals are expected to breed earlier in response to temperature cues. However, if species use fixed cues such as daylight, their breeding timing might remain fixed, potentially creating disconnects between their life history and environmental conditions. Wood frogs Rana sylvatica are a cold-adapted species that reproduce in early spring, immediately after breeding ponds are free of ice. We used long-term surveys of wood frog oviposition timing in 64 breeding ponds over 20 yr to show that, despite experiencing a warming of 0.29°C per decade in annual temperature, wood frog breeding phenology has shifted later by 2.8 d since 2000 (1.4 d per decade; 4.8 d per °C). This counterintuitive pattern is likely the result of changes in the timing of snowpack accumulation and melting. Finally, we used relationships between climate and oviposition between 2000 and 2018 to hindcast oviposition dates from climate records to model longer-term trends since 1980. Our study indicates that species can respond to fine-grained seasonal climate heterogeneity within years that is not apparent or counterintuitive when related to annual trends across years.     

Diversity of butterflies on British islands: ecological influences underlying the roles of area, isolation and the size of the faunal source

To distinguish between the influences of area and isolation on the butterfly faunas of British islands two approaches are adopted. First, species richness is related to island area, isolation and the size of the faunal source. Neither area nor isolation account for much variance in species richness, though area is more important than isolation. In contrast, species richness corresponds closely to the size of the faunal source on nearby islands and to that at proximate locations on adjacent mainlands. The second approach relates the incidence of species on islands to their ecological attributes. A very close relationship is found between species incidence on islands and those ecological variables that measure potential for migration and colonization and that resist extinction. The implications are that the majority of British islands in this survey are insufficiently isolated to prevent intermittent migrations of butterflies to them or so small as to generate frequent extinctions. Independent data indicate the capacity of many resident species to migrate distances in excess of the isolation of most of the islands. Some evidence also exists for the long-term survival of species on islands; important considerations in this respect are that most islands in the survey are large compared to habitat patches sustaining species on mainland Britain and that substantial portions of islands are retained in early seral stages or comprise long-lived stable habitats (e.g. peat mosses) that are particularly suitable for many British species.     

Long-term trends in the arrival dates of spring migrants

Data on the spring arrival dates of 23 species of migrants in Leicestershire over a 50-year period are presented. Chiffchaff, Sand Martin, Blackcap and Sedge Warbler showed a significant trend towards earlier arrival over the period, while Tree Pipit, Cuckoo, Whinchat, Whitethroat and Garden Warbler showed a significant trend towards later arrival. Fifteen species arrived noticeably earlier in the 1940s, a period of warm springs, while several species showed earlier arrivals in the 1980s. A number of species showed later arrival dates in the 1960s and 1970s, when April temperatures were colder than average. Several species showed significant correlations between arrival date and temperature. Arrival dates of the earliest species were much more variable than those arriving later, while species arriving in the second half of April showed a generally synchronous arrival. The results are discussed in the context of global warming.     

Habitat associations of thermophilous butterflies are reduced despite climatic warming

Climate warming threatens the survival of species at their warm, trailing‐edge range boundaries but also provides opportunities for the ecological release of populations at the cool, leading edges of their distributions. Thus, as the climate warms, leading‐edge populations are expected to utilize an increased range of habitat types, leading to larger population sizes and range expansion. Here, we test the hypothesis that the habitat associations of British butterflies have expanded over three decades of climate warming. We characterize the habitat breadth of 27 southerly distributed species from 77 monitoring transects between 1977 and 2007 by considering changes in densities of butterflies across 11 habitat types. Contrary to expectation, we find that 20 of 27 (74%) butterfly species showed long‐term contractions in their habitat associations, despite some short‐term expansions in habitat breadth in warmer‐than‐usual years. Thus, we conclude that climatic warming has ameliorated habitat contractions caused by other environmental drivers to some extent, but that habitat degradation continues to be a major driver of reductions in habitat breadth and population density of butterflies.     

Living on the edge: population ecology of <Emphasis Type="Italic">Phengaris teleius</Emphasis> in Serbia

Due to their complex and interesting life cycle and alarming conservation status the Large Blue butterflies have become one of the most studied group of insects in Europe. In Serbia, however, Phengaris teleius has recently been discovered (in the far north of the country) and, since this initial finding, significant efforts have been made to map the local distribution of this species and to implement conservation measures. A mark-release-recapture study was initiated to obtain a more detailed report about population size and structure in Serbia. Results have shown that localities and patches within these localities are well connected by migration of butterflies and gave some evidence for metapopulation organisation. The total number of individuals was estimated at 15,000, which makes it one of the largest known metapopulations in Europe. Unusual for populations at the edge of the distribution range, these are characterized with large population estimates and relative stability. Compared to other areas in Europe, butterflies in Serbia start to fly 2 weeks later in the summer, probably a consequence of a late mowing regime initiated during July. Some evidence of a negative relationship between the survival of P. teleius adults and local population density was also found. With favourable management, these populations could remain stable for the foreseeable future. The preservation of large, healthy populations of P. teleius supported by a favourable management and monitoring programme along with the sustainability of individual dispersal translocation between patches should be a prime aim.     

A multivariate approach to the determination of faunal structures among European butterfly species (Lepidoptera: Rhopalocera)

Multivariate analyses of 393 butterfly species over 85 geographical areas (R- and Q-data matrices) in Europe and North Africa have produced a consistent pattern of faunal structures (units and regions). Prominent features to emerge are the latitudinal zonation of geographical units and the division of the Mediterranean into western and eastern components; southwards in Europe, endemicity increases whereas faunal structures decrease in spatial dimensions. Central Europe–from the Urals to the British Isles–forms a single large faunal structure (extent unit and region). A model has been constructed to account for Pleistocene evolutionary changes and endemism in European butterflies and for the east-west taxonomic divisions in the extent faunal structure which dominates central Europe. Periodic Pleistocene climatic changes have resulted in cycles of population extinction, isolation, evolution and migration, but the nature and timing of events has depended on the environmental tolerances of species belonging to different faunal units. During Pleistocene glaciations, southern species have been relatively static and more isolated and have evolved independently. By comparison, northern species have been more mobile and have migrated over large distances. Contact and hybrid zones among cosmopolitan species in northern Europe are probably of some antiquity. They result from persistent survival and isolation of refuge populations in the west and east Mediterranean during glacial phases; dispersal from these refuges leads to their regeneration during each interglacial.