Larval survival in populations of the large copper butterfly Lycaena dispar batavus

Lycaena dispar is recorded as an early case of extinction and subsequent re-introduction in the UK. However, repeated establishment attempts have not resulted in self-sustaining populations, including those having taken place at Woodwalton Fen (Cambridgeshire, UK).
Earlier studies at Woodwalton Fen showed that the highest mortalities occurred between the egg stage and resumption of larval feeding in spring.
This study was designed; firstly, to investigate factors causing field mortality during different larval stages; secondly, to compare survival on Rumex hydrolapathum foodplants in different habitat situations; and thirdly, to compare survival in a natural population (Weerribben, The Netherlands) with the introduced Woodwalton Fen population. Experiments employed exclusion cages in order to examine the relative roles of vertebrate and invertebrate natural enemies, and survival on food-plants in open fen and waterside situations was compared.
Results suggest invertebrate predation to be the dominant mortality factor acting upon pre-diapause larvae. Over the winter diapause natural enemies do not have a significant role. However during both these stages losses still occur that are unaccounted for. Extensive winter flooding appears to increase overwintering losses. Vertebrate predators cause significant mortality of post-diapause larvae.
No significant difference was found between larval survival on open fen versus waterside plants.
Comparison of introduced and native populations reveal that overwintering survival was significantly higher in the latter; potential reasons for this difference are discussed.     

Habitat requirements of Lycaena dispar batavus and implications for re-establishment in England

This paper reports on studies of the habitat requirements of Lycaena dispar batavus in its largest remaining site, the Weerribben National Park in the Netherlands. Observations and experiments were carried out to assess requirements for male territories, egg-laying sites and larval survival. The management of the fenland was also assessed on a landscape scale. Male territories were all situated in summer cut fen meadow. Eggs were laid on Rumex hydrolapathum in a range of habitat situations, although a preference was shown for plants on fen edges. Larval survival was highest on plants on watersides and fen edges, with no survival on plants in summer cut fen meadow. The findings are discussed in relation to whether the Broadland area in eastern England is suitable as a re-establishment site. Although many elements of the habitat mosaic are present, at the landscape scale there is insufficient open fenland and areas suitable for male territories are probably too isolated; however, a fen restoration strategy is in place which may restore suitable habitat and make re-establishment feasible in the future.     

Phylogeography, genetic diversity and conservation of the large copper butterfly Lycaena dispar in Europe

We argue that insect species conservation at large scales should take account of the distribution of genetic diversity among populations. Maintenance of genetic diversity may be vital in retaining a species' adaptive capacity and evolutionary potential. We illustrate the concept using the example of the large copper butterfly Lycaena dispar in Europe. This species has become extinct in parts of its range and is declining rapidly in others, whilst conversely, increasing in many areas. The latter has recently reduced its conservation status. Mitochondrial DNA analysis is used to construct a phylogeography from a preliminary sample set obtained from across Europe. A cytochrome b fragment of 402 base pairs was sequenced and 10 haplotypes were found. Relatedness among populations suggest that those from northern and central Europe are closely related and probably form one evolutionary significant unit (ESU) reflecting post-glacial colonization from southeast Europe. In contrast, the sample from Italy is divergent and should be considered a separate ESU. Our results, combined with ecological data, suggest that conservation action for this species should be targeted on specific regions and populations.     

Effects of mire type, land use and climate on a strongly declining wetland butterfly

Mires are characterised by highly specialised and threatened wildlife. One of these mire specialists that has severely declined is the Large Heath (Coenonympha tullia). However, detailed studies concerning the reasons for the dramatic population loss in central Europe are missing so far. In this paper: (1) we analyse the habitat preferences of adult C. tullia and oviposition site selection with respect to mire type and land use in one of the last German strongholds, and (2) we attempt to disentangle the impacts of land use and climate change on the decline of C. tullia on a national scale across Germany. Abundance of adult C. tullia was clearly affected by mire type and land use. It was highest on managed calcareous fens and lowest on unmanaged transition and raised bogs. The oviposition pattern of C. tullia females was best explained by (1) quantity of potential host plants (Eriophorum latifolium), (2) vegetation structure, and (3) microclimate. In Germany, C. tullia populations have become extinct in lowlands with a mild and relatively dry climate while most colonies in wet and cold mountain areas have survived. However, there is evidence that habitat loss and habitat deterioration, rather than climate change, are the drivers of the range retraction. To create low-growing vegetation rich in E. latifolium, traditional mowing late in the year and light grazing seem to be suitable management tools. In addition, conservation management should aim to maintain high water levels or restore them, especially to counteract effects of climate change in the future.     

Reinstatement of a New Zealand copper butterfly,Lycaena rauparaha (Fereday, 1877)

Abstract

Previous confusion over the nomenclature and status of the New Zealand copper butterflies is reviewed. Lycaena rauparaha n. comb., which has been regarded as a variety of L. salustius in recent literature, is reinstated and adult, larval, and pupal characters are described. Its distinctions from L. salustius and L. feredayi are tabulated and a key to this species complex is given.     

Environmental induction of larval diapause and life-history consequences of post-diapause development in the Large Copper butterfly,Lycaena dispar (Lepidoptera: Lycaenidae)

Many insects in temperate zones withstand the adverse conditions of winter through entering diapause and the two most important environmental stimuli that induce diapause are photoperiod and ambient temperature. The Large Copper butterfly, Lycaena dispar Haworth (Lepidoptera: Lycaenidae), is a Palearctic butterfly that hibernates as larvae. Since this butterfly is a near threatened species in some regions, there has been a growing need for a standardized protocol for mass rearing of this butterfly based on the adequate knowledge of its ecology. In the present study, we first identified that L. dispar larvae were sensitive to the photoperiodic induction of diapause during their first larval instar. We then investigated to what extent the diapause-inducing effects of photoperiod could be modified by ambient temperatures in L. dispar larvae by exposing them to the range of day-lengths (L:D 14:10, 12:12, 10:14 and 8:16) at three different temperatures (15, 20 and 25 °C). All larvae were induced to enter diapause at low ambient temperature (15 °C) regardless of photoperiod, whereas most of them (86 %) exhibited direct development when temperature was high (25 °C). The photoperiodic induction of diapause was evident when day-length was shorter than 14 h at intermediate temperature (20 °C). Pre-diapause development was prolonged at low temperatures. Finally, we found that post-diapause development of L. dispar larvae was determined by both the chilling temperature experienced by diapausing larvae and the duration of larval diapause. Adult emergence was enhanced when larvae were chilled at 8 °C and when they had been under the state of diapause for 20 days before they were treated to terminate diapause.     

The distribution and decline of a widespread butterfly Lycaena phlaeas in a pastoral landscape

1. Ecological specialists are often regarded as most likely to be threatened by anthropogenic habitat changes but few relevant data are available on changes in the status of widespread species. 2. Grid square distribution maps have been used widely to measure rates of decline and target conservation resources but it is known that coarse grain mapping is not appropriate to identify declines in widespread species that initially contain numerous local populations per grid cell. Changes in the status of widespread species need to be quantified. 3. Present‐day habitat associations, determined from over 2000 transect counts, combined with data on historical and present‐day habitat distributions, reveal that the area of occupancy and population‐level rate of decline of the Small Copper butterfly Lycaena phlaeas is likely to have been of the order of 92 and 89% respectively, in 35 km² of North Wales. Similar data on the species' major host plants Rumex acetosa and R. acetosella indicate possible declines in area occupied of 48 and 91%. If a 1‐km² grid was applied to the landscape, and if L. phlaeas, R. acetosa, and R. acetosella had occupied all 1‐km² cells in the study area in 1901 (non‐limestone cells for R. acetosella only), their declines would only have been recorded as 15, 9, and 35% respectively. 4. Many declining ecological specialists are threatened with extinction because of their initial rarity. At a population level, however, they may or may not be declining faster than less specialised species. The results presented here illustrate that some widespread species may have declined as much as many of Britain's rarities.     

春甘蓝田菜青虫虫情简便监测技术

在湖南 ,春甘蓝的大田生长期一般在 1月下旬至 6月上旬之间 ,这一季节的主要食叶害虫是菜青虫 Pierisrapae L .,其次是灯蛾 ,过去大量发生的小菜蛾近年来一直处于低数量低水平。而甜菜夜蛾、斜纹夜蛾此时发生极少。灯蛾的卵呈块产 ,卵粒裸露 ,单层排列 ,低龄幼虫呈集团性分布 ,不难发现。为此 ,我们只针对主要害虫菜青虫作了虫情简便监测技术的研究。1　方法1 .1　于 1 998年 5月 3日至 6月 4日菜青虫主要发生危害期在甘蓝地及其周边共 0 .0 5 km2的固定范围内每日上午 1 0时计数飞舞的成虫(菜粉蝶 )数 ,一般计数 3～ 5次 ,求其平均值 ;并…     

Contrasting patterns of wetland use by sympatric larids in winter

The population dynamics and behaviour of the larid assemblage of a Mediterranean coastal wetland, the Vourkari inlet in Greece, were studied during the winter of 2008–2009. More black-headed gulls (Larus ridibundus) were seen in the inlet in December, more Mediterranean gulls (Larus melanocephalus) were present from mid-January to mid-February, while little variation was observed in yellow-legged gull (Larus michahellis) numbers throughout winter. Bird numbers remained stable through the day for the yellow-legged gull, but fewer black-headed and Mediterranean gulls were present in the late morning than the other periods. Diurnal activity patterns showed that feeding was the primary behaviour for yellow-legged gulls and resting and sleeping for black-headed and Mediterranean gulls, with locomotion being equally important for all species. All larids were primarily feeding in the late morning period, but yellow-legged gulls were doing so in much higher proportions. These contrasting patterns suggested that the Vourkari inlet was more important as a day roost for black-headed and Mediterranean gulls and a preferred feeding ground for yellow-legged gulls. Furthermore, results suggested that resting and sleeping were interchangeable activities and all other activities had more or less the same time demands on a daily basis, and also a consistent pattern across species in sleeping proportions that might indicate cross-species synchrony in sleep patterns; however future research is needed to resolve these issues. This study provided important new information on the winter ecology of three larids and revealed patterns of wetland use by these species that could help assess the importance of certain areas and improve coastal habitat management strategies to benefit birds.     

Patch occupancy in the endangered butterfly Lycaena helle in a fragmented landscape: effects of habitat quality, patch size and isolation

While there is agreement that both habitat quality and habitat network characteristics (such as patch size and isolation) contribute to the occupancy of patches by any given species, the relative importance of these factors is under debate. This issue is of fundamental ecological importance, and moreover of special concern for conservation biologists aiming at preserving endangered species. Against this background we investigated patch occupancy in the violet copper Lycaena helle, one of the rarest butterfly species in Central Europe, in the Westerwald area (Rhineland-Palatinate, Western Germany). Occupied (n = 102) differed from vacant (n = 128) patches in altitude, size, connectivity, availability of wind shelter, in the abundance of the larval host-plant, in the abundance of a grass species indicating favorable habitat conditions and in the abundance of nitrophilous plants. Overall, patch occupancy was primarily determined by patch size, connectivity and the abundance of the larval host plant, while all other parameters of habitat quality were of subordinate importance. Therefore, our findings suggest that even for extremely sedentary species such as L. helle habitat networks are decisive and—next to the preservation of habitat quality—need to be an integral part of any conservation management for this species.     

Impact of grazing management on hibernating caterpillars of the butterfly Melitaea cinxia in calcareous grasslands

Semi-natural grasslands are increasingly grazed by large herbivores for nature conservation purposes. For many insects such grazing is essential for the conservation of their habitat, but at the same time, populations decrease at high grazing intensity. We hypothesised that grazing management may cause increased butterfly mortality, especially for life-stages with low mobility, such as hibernating caterpillars. To test this, we measured the effect of sheep grazing on overwinter larval survival. We used the Glanville fritillary (Melitaea cinxia), which has gregarious caterpillars hibernating in silk nests, as a model species. Caterpillar nests were monitored throughout the hibernating period in calcareous grassland reserves with low and high intensity sheep grazing and in an ungrazed control treatment. After grazing, 64?% of the nests at the high intensity grazing treatment were damaged or missing, compared to 8 and 12?% at the ungrazed and low intensity grazing treatment, respectively. Nest volume and caterpillar survival were 50?% lower at the high intensity grazing treatment compared to both ungrazed and low intensity grazing treatments. Nest damage and increased mortality were mainly caused by incidental ingestion of the caterpillars by the sheep. It is likely that grazing similarly affects other invertebrates, depending on their location within the vegetation and their ability to actively avoid herbivores. This implies that the impact of grazing strongly depends on the timing of this management in relation to the phenology of the species. A greater focus on immature and inactive life-stages in conservation policy in general and particularly in action plans for endangered species is required to effectively preserve invertebrate diversity.     

Impact of dietary allelochemicals on gypsy moth (Lymantria dispar) caterpillars: importance of midgut alkalinity

Midgut pH of gypsy moth larvae was depressed artificially with buffered diet to examine the impact of alkalinity on the caterpillars' ability to tolerate a dietary polyphenol and a quinone. A 2x3 factorial design was used, with 2 levels of succinate buffer and 3 dietary amendments (tannic acid, juglone, or control). Development was monitored during the third and fourth instars, with consumption, food passage rates, midgut pH, and midgut redox potential (Eh) measured in the fourth instar. Diet buffering successfully depressed midgut pH to hypothetically suboptimal acidic levels without reductions in survivorship, but it did reduce larval growth and impede development. Buffering dramatically reduced survivorship of fourth instar larvae eating diets containing tannic acid or juglone. Growth increased on unbuffered diet amended with tannic acid, but not with juglone. Caterpillars passed food through the gut more slowly when feeding on buffered tannic acid diet or on unbuffered juglone diet. These results indicate that maintenance of midgut alkalinity is critical to tolerance of dietary tannic acid and juglone, and that these allelochemicals have very different activities in the caterpillar gut.     

Photoperiod- and temperature-dependent regulation of pupal beige/black polymorphism in the small copper butterfly, Lycaena phlaeas daimio Seitz

The small copper butterfly, Lycaena phlaeas daimio, has pupal beige/black polymorphism, the development of which is found to be controlled in an apparent association with the development of adult seasonal polymorphism (spring and summer morphs) by photoperiod and temperature in the larval stages. That is, the pupae of beige and black types developed under long-day and short-day conditions tend to develop into brown-winged and red-winged adults, respectively. In addition, a large proportion of long-day pharate pupae chilled at 4 degrees C for 5 days were observed to develop into pupae whose head-thoracic complexes and abdomens were judged to be of the black and intermediate types, respectively. They developed into adults with redder wings as compared to those obtained from unchilled pupae. The results indicate that the physiological mechanism underlying the photoperiodic control of the development of adult seasonal polymorphism may also play a significant role in the determination of pupal beige/black polymorphism in L. phlaeas daimio. Furthermore, cuticle melanization was found to be induced in the head-thoracic complexes of pupae by chilling of the pharate pupae. Melanization of pupal cuticle seems to occur in a close association with the development of reddish-winged adults.     

Habitat utilisation by pike Esox lucius L. during winter floods in a southern English chalk river

Seven pike Esox lucius L., implanted with radio-transmitters, were tracked throughout autumn and winter in the River Frome, a southern English chalk river. During the first flood events of the year, pike remained within the main river channel but during subsequent flood events, pike could also be found in flooded fields, in drainage ditches or in a millstream. Eighty percent of the fixes over flooded land occurred within 10 m of the riverbank, although distances of up to 89 m from the bank were recorded. In ditches, pike could be found over 250 m from the main river. For pike in ditches and flooded fields, distance from the main river channel was positively correlated with discharge. There was individual variation amongst pike for the habitat types selected, with some pike utilising flooded field or ditch habitat more often than others. The proportion of time spent out of the main river channel does not appear to be related to the size of the pike. It is hypothesised that pike are leaving the main river channel to exploit feeding opportunities in the flooded fields and drainage ditches, rather than using these areas as refugia from high flow conditions.     

Effects of flooding on a wetland bird community

The Lake Izunuma–Uchinuma was extensively flooded in 1998 and vast areas (approximately 200 ha) of rice fields were submerged for the first time since the land was reclaimed in 1963. When the numbers of birds in 1998 were compared with those in normal years (1995–1997), the numbers of black-crowned night herons (Nycticorax nycticorax L.), cattle egrets (Bubulcus ibis L.) and black kites (Milvus migrans Boddaert) increased during the flooding, and the numbers of ducks, especially dabbling ducks (seven species of 11 Anas spp.) were high even after the flooding, while the numbers of the little grebe Tachybaputus ruficollis Pallas were diminished by the flooding.     

Habitat distributional change of the large copper,Lycaena dispar (Lepidoptera: Lycaenidae), on the Korean Peninsula: Studies along the Geum River

Habitat changes of butterflies are caused by various factors, including food plants, natural enemies, and climate change. Recently, new habitats of the large copper have been reported on the Korean Peninsula, mostly along a riverside. This implies that this butterfly may move along a drainage system. In this study, new habitats of the large copper along the Geum River were investigated and the population dynamics of the butterfly were monitored in selected areas through the mark–release–recapture method. The results showed that the large copper had dispersed inland along the Geum River, which runs along the central western seashore of South Korea. In addition, the habitat distribution of the butterfly showed good coincidence with the distribution of a sorrel, Rumex japonicas, the main food plant of the large copper. A study on the moving distance of the butterfly showed that females moved further than males, while males showed more frequent movement within a short distance. The movement of the large copper along the Geum River, which lies over 100 km south of the Han River (a habitat previously reported in 1973), suggests that the movement of the butterfly might be more affected by human activities, such as the development of bike paths along the Geum River and maintenance of the riverbed, than climate change.     

Conserving a wetland butterfly: quantifying early lifestage survival through seasonal flooding, adult nectar, and habitat preference

Conservation of once thought extinct populations of Lycaena xanthoides in western Oregon will require specific information of how the butterfly interacts with its now rare wetland habitat. Three experiments were conducted to yield information directly applicable to wetland restoration work: (1) to quantify the survival of L. xanthoides eggs laid on inundated plants, (2) to quantify adult nectar preferences, (3) to investigate the role of adult resources and how butterflies assess habitat quality. Survival of eggs laid on inundated plants was nearly seven times lower than the survival of eggs laid on uninundated plants, indicating that eggs laid on plants that are seasonally flooded are a population sink. Adult L. xanthoides preferred an endemic native nectar plant, Grindelia integrifolia × nana, as an adult resource and used it approximately 88% of the time while other butterfly species preferred to nectar on the non-native Mentha pulegium. Adult L. xanthoides had a significantly greater short-term recapture rate in two restored study sites that had a high relative amount of Grindelia integrifolia × nana compared to a degraded site that lacked the preferred nectar source. Based on the results from the three small experiments, restoration of wetlands for L. xanthoides should concentrate plantings of host plant in non-flooded areas and propagate conspicuous patches of the preferred nectar plant.     

Effects of cytoplasmic polyhedrosis virus on diapausing Heliothis virescens

Laboratory studies were conducted to determine effects of cytoplasmic polyhedrosis virus on diapausing Heliothis virescens. Most virus-infected individuals died in the larval stage. Infected pupae yielded as many moths as healthy. Females from surviving virus-infected larvae produced fewer eggs than those from healthy larvae, but there was no statistical difference in longevity of adults between healthy and infected groups. Infected moths yielded lower than normal quantities of extracted fatty acids.     

Impact of predation by a cave-dwelling bat, Rhinolophus ferrumequinum, on the diapausing population of a troglophilic moth, Goniocraspidum preyeri

The predation pressure of the greater horseshoe bat, Rhinolophus ferrumequinum, on the diapausing population of the noctuid moth, Goniocraspidum preyeri, was examined at an abandoned mine in central Japan. These bats did not prey on the moths in summer. The number of moths preyed on was largest in March. More than 60% of the diapausing moths were eaten by only three or four bats, which accounted for over 90% of all the moth deaths. This moth was an important source of food at the end of the hibernating period when the bat had already used most of its stored subcutaneous fat. The predation on G. preyeri may have raised the overwintering success rate of R. ferrumequinum.