Maculinea Nausithous Research Articles

What limits the spread of two congeneric butterfly species after their reintroduction: quality or spatial arrangement of habitat?

AbstractPopulation growth and spread of recently reintroduced species is crucial for the success of their reintroduction. We analysed what limits the spread of two congeneric butterfly species Maculinea teleius and Maculinea nausithous, over 10 years following their reintroduction. During this time, their distributions appeared to be limited to a few sites although it was thought that more suitable habitats were available. Thus, we question, does the quality or the spatial arrangement of their habitat limit their spread? Although adult individuals of both species can select high‐quality plots, we show that selection of suitable plots in the area of reintroduction is spatially constrained. A low colonization probability of unoccupied distant plots of high quality was found for both species. The abandonment of occupied plots in Ma. teleius was also found to be dependent on the distance to occupied plots. We conclude that the spatial distribution of the two species during the 10 years following reintroduction was limited by the spatial arrangement of their habitat, rather than by the availability of high‐quality plots. The spatial constraints in movement can explain observed source–sink structures when female butterflies deposit their eggs on low‐quality plots. We conclude that although these species have very similar life histories, they require different approaches to their conservation due to subtle differences in adult habitat use and movement. Conservation of Ma. teleius should concentrate on improving local habitat quality, whereas conservation of Ma. nausithous is predicted to be more effective by creating a spatial network of suitable habitat plots, such as along road verges.

Habitat requirements and niche selection of Maculinea nausithous and M. teleius (Lepidoptera: Lycaenidae) within a large sympatric metapopulation

Identifying the factors determining the distribution and abundance of species is a fundamental question in population ecology. Based on habitat characteristics, we sought to identify factors affecting the abundance of two vulnerable large blue butterflies, Maculinea teleius and Maculinea nausithous. Both species often occur in sympatry and thus seem to have very similar requirements with regard to their wetland habitats. Even though, M. nausithous is usually far more widespread and abundant than M. teleius in Central Europe. For understanding this difference despite a similar ecology, more data is needed on the niche selection of both species. We have recorded Maculinea numbers, vegetation structure, food-plant abundance and ant community for 120 patches within a large sympatric metapopulation of both species in the Westerwald area (Germany). Unexpectedly (based on previous data), both species were equally common in the study area, reaching high densities of up to 61 (M. teleius) and 58 (M. nausithous) individuals per 50 m². In 116 out of 120 patches both species occurred sympatrically. The preferred habitat was extensively managed grassland, being mown or grazed twice a year. The abundance of both Maculinea species was positively related to vegetation height, food-plant and Myrmica rubra abundance. There was no association between M. teleius and its putative main host-ant Myrmica scabrinodis, indicating that M. rubra may serve as an important host in the Westerwald area. The positive effect of food-plant abundance was more pronounced for M. teleius, coinciding with a higher food-plant abundance at sites dominated by M. teleius compared to those dominated by M. nausithous. Thus, M. teleius seems to be slightly more demanding than M. nausithous in terms of food-plant availability.

Butterflies (Lepidoptera: Hesperioidea, Papilionoidea) of the Kampinos National Park and its buffer zone

Poland; e-mail: marcins@uwb.edu.pl Abstract: Kampinos National Park is the second largest protected area in Poland and therefore a potentially important stronghold for biodiversity in the Mazovia region. However it has been abandoned as an area of lepidopterological studies for a long time. A total number of 80 butterfly species were recorded during inventory studies (2005–2008), which proved the occurrence of 80 species (81.6% of species recorded in the Mazovia voivodeship and about half of Polish fauna), including 7 from the European Red Data Book and 15 from the national red list (8 protected by law). Several xerothermophilous species have probably become extinct in the last few decades (Colias myrmidone, Pseudophilotes vicrama, Melitaea aurelia, Hipparchia statilinus, H. alcyone ), or are endangered in the KNP and in the region (e.g. Maculinea arion, Melitaea didyma), due to afforestation and spontaneous succession. Higrophilous butterflies have generally suffered less from recent changes in land use, but action to stop the deterioration of their habitats is urgently needed. Lycaena dispar, Maculinea teleius and M. alcon are still quite widespread but L. helle and Euphydras aurinia were recorded on single sites only. However, Maculinea nausithous was observed only in 2005 and has probably just disappeared from the KNP. Despite the aforementioned losses, the Kampinos Forest deserves to be added to the list of the Prime Butterfly Areas in Europe. Key words: butterflies, biodiversity, Kampinos National Park, fauna of Poland, endangered species

Adopting the Precautionary Principle in Designing and Managing Natura 2000 Areas (Exemplified by the Conservation of the Butterfly Maculinea Nausithous in a Rural Landscape North of Dresden (Saxony))

Adopting the Precautionary Principle in Designing and Managing Natura 2000 Areas (Exemplified by the Conservation of the Butterfly Maculinea Nausithous in a Rural Landscape North of Dresden (Saxony)) The precautionary principle is more and more incorporated into national law and decision-making on natural resource management and biodiversity conservation. In the coherent European network of protected areas Natura 2000, the precautionary principle finds expression in the obligation to provide favourable conditions for the long-term survival of species and habitats, especially of the priority ones listed in the annexes of the Habitats Directive and the Birds Directive. After describing principles, structure, implementation and procedures of this rather new instrument for nature conservation using the example of one of the various Natura 2000 areas in Saxony (Germany), opportunities and problems for biodiversity conservation are outlined with particular regard for the situation in an agricultural landscape. Special attention is given to the following questions: requirements of and actual threats to the target species (the butterfly Maculinea nausithous), legal means and economic incentives for suitable measures, the management plan, and the role of stakeholders. It turns out that Natura 2000 could be an effective tool to advance nature conservation, and with special regard to the precautionary principle. Every effort is necessary to gain more public acceptance of Natura 2000, as well as to improve scientific knowledge concerning species and habitats under protection.

Populations with Explicit Borders in Space and Time: Concept, Terminology, and Estimation of Characteristic Parameters

Biologists studying short-lived organisms have become aware of the need to recognize an explicit temporal extend of a population over a considerable time. In this article we outline the concept and the realm of populations with explicit spatial and temporary boundaries. We call such populations "temporally bounded populations". In the concept, time is of the same importance as space in terms of a dimension to which a population is restricted. Two parameters not available for populations that are only spatially defined characterise temporally bounded populations: total population size, which is the total number of individuals present within the temporal borders, and total residence time, which is the sum of the residence times of all individuals. We briefly review methods to estimate these parameters. We illustrate the concept for the large blue butterfly (Maculinea nausithous) and outline insights into ecological and conservation-relevant processes that cannot be gained without the use of the concept.

Population structure of a large blue butterfly and its specialist parasitoid in a fragmented landscape

Habitat fragmentation may interrupt trophic interactions if herbivores and their specific parasitoids respond differently to decreasing connectivity of populations. Theoretical models predict that species at higher trophic levels are more negatively affected by isolation than lower trophic level species. By combining ecological data with genetic information from microsatellite markers we tested this hypothesis on the butterfly Maculinea nausithous and its specialist hymenopteran parasitoid Neotypus melanocephalus. We assessed the susceptibility of both species to habitat fragmentation by measuring population density, rate of parasitism, overall genetic differentiation (theta(ST)) and allelic richness in a large metapopulation. We also simulated the dynamics of genetic differentiation among local populations to asses the relative effects of migration rate, population size, and haplodiploid (parasitoid) and diploid (host) inheritance on metapopulation persistence. We show that parasitism by N. melanocephalus is less frequent at larger distances to the nearest neighbouring population of M. nausithous hosts, but that host density itself is not affected by isolation. Allelic richness was independent of isolation, but the mean genetic differentiation among local parasitoid populations increased with the distance between these populations. Overall, genetic differentiation in the parasitoid wasp was much greater than in the butterfly host and our simulations indicate that this difference is due to a combination of haplodiploidy and small local population sizes. Our results thus support the hypothesis that Neotypus parasitoid wasps are more sensitive to habitat fragmentation than their Maculinea butterfly hosts.

Genetic differentiation among the Maculinea species (Lepidoptera: Lycaenidae) in eastern Central Europe

The present study aimed to analyse the level of genetic variation in the eastern Central European (Slovenia, Hungary, and Romania/Transylvania) populations of the Large Blues (Maculinea) to analyse the pattern of differentiation both between and within the species. One objective was to compare the level of differentiation between the two disputed species (Maculinea alcon and Maculinea rebeli) with that among the other species. Imagos were collected from 23 localities in eastern Central Europe in 2002. Enzyme polymorphism was analysed using polyacrylamide gel electrophoresis. Fourteen enzyme loci were studied in all samples. In the analysis of the data, F-statistics and Nei’s genetic distances were calculated and a dendrogram (unweighted pair group method with arithmetic mean) was constructed on the basis of the distance matrix. A multivariate analysis of variance was performed to study the pattern of genetic differentiation among the samples. Principal component analysis analysis was also carried out using the allele frequency data of the samples. Our results indicated that the large blues are generally less polymorphic than other European lycaenid butterflies studied. At the same time, the level of genetic differentiation was high, even among local populations within the species. A low level of genetic variation within the populations coupled with strong differentiation among them implies the effect of genetic drift. Strong genetic differentiation of four Maculinea species (M. alcon, Maculinea teleius, Maculinea nausithous, and Maculinea arion) was confirmed. Significant differentiation was not found between M. alcon and M. rebeli. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91, 11–21.

Effects of management cessation on grassland butterflies in southern Poland

The effects of natural succession and invasions of indigenous reed Phragmites australis and alien goldenrods Solidago sp. on butterfly communities of wet grassland were investigated in Kraków (southern Poland) after management cessation. The total number of species and individuals, mean number of species per survey and a species diversity (Shannon-Wiener index) were highest in fallow lands, old fallow lands and young forests, and the lowest in fallow lands invaded by reed and goldenrod and in mature forests. These results (1) reveal the importance of extensively mown meadows and fallow land for butterflies, (2) indicate that meadow restoration is still possible even several decades after abandonment and (3) show that invasion of reed and non-native goldenrods should be prevented. A few very rare species included in the Habitat Directive ( Lycaena dispar, Lycaena helle, Maculinea teleius and Maculinea nausithous) were common on the wet grasslands in Krakow, thus the area should be protected. Moreover, it was shown that the presence and relative abundance of Maculinea butterflies were good indicators of general butterfly species richness and abundance in the investigated wet grasslands.

Spatial patterns of host exploitation in a larval parasitoid of the predatory dusky large blue Maculinea nausithous

Summary The foraging behaviour of the parasitoid wasp Neotypus melanocephalus and factors affecting parasitism at the population level were studied. This specialised parasitoid attacks caterpillars of the butterfly Maculinea nausithous , which sequentially feed on the plant Sanguisorba officinalis and specific red Myrmica ants. Among M. nausithous populations, there is considerable variation in caterpillar densities. At low M. nausithous densities, foraging might be time consuming for N. melanocephalus. High host densities may not always be advantageous to foraging parasitoids due to the caterpillars’ frequent overexploitation of ant resources and subsequent density-dependent mortality. In order to disperse progeny, we hypothesised that N. melanocephalus should search in a non-random way at the level of the micro-habitat, i.e., single flower heads of S. officinalis . Our analysis of 32 natural populations in the Upper Rhine valley in Germany did not show a density-dependent relationship between M. nausithous caterpillars and parasitism. Furthermore, habitat parameters like patch size and density of the host's food plant did not affect the parasitism rate. Foraging N. melanocephalus females preferred to search on large flower heads. They probed host-occupied flower heads only, visiting non-host-exploited flower heads only briefly. Time spent on a flower head was independent of the number of caterpillars per flower head. This study indicates that N. melanocephalus increases its foraging efficiency by preferring large flower heads that were previously shown to contain more host caterpillars than small flower heads. Furthermore, oviposition increases the likelihood of continuing to search on a flower head, which is an adaptive strategy for parasitoids foraging for aggregated hosts. However, many host-occupied flower heads were not probed by N. melanocephalus . We discuss the possibility that temporal host refuges of M. nausithous caterpillars might contribute to heterogeneity of parasitism, and why spreading offspring might constitute a suitable strategy for a parasitoid of an ant-parasitic butterfly.

Nine polymorphic microsatellite loci for the parasitic wasp Neotypus melanocephalus (Hymenoptera: Ichneumonidae)

AbstractNine polymorphic microsatellite loci were isolated from Neotypus melanocephalus (Gmelin), a parasitoid of the parasitic large blue butterfly Maculinea nausithous. Allelic diversity and heterozygosity were quantified in samples from the Upper Rhine valley in Southwest Germany.

Influence of mowing on the persistence of two endangered large blue butterfly species

Summary Mowing influences two endangered butterfly species, Maculinea nausithous and Maculinea teleius, directly through egg destruction and larval mortality on the mown plants and indirectly through altering the abundance of their sequential resources in meadows (Sanguisorba plants for oviposition and early larval development and Myrmica ant nests for later larval development and pupation). Although conservation biologists have argued that mowing during the adult stage is detrimental to population persistence, it is not obvious how the timing and frequency of mowing impact on population dynamics. A simulation model was used to investigate how current ‘traditional’ mowing regimes could be altered to reconcile butterfly conservation with agriculture. The key mechanism affecting the impact of mowing on population persistence was the interaction between density‐independent and density‐dependent mortalities in different larval stages of each life cycle. Because of this interaction, optimal mowing regimes for butterfly conservation were sensitive to the type of density regulation displayed by each species, and to landscape attributes such as the influence of climate on resource availability and the level of parasitism. Despite this sensitivity, we were able to identify robust mowing regimes appropriate for a wide range of landscape attributes and to derive general management recommendations. Synthesis and applications. Our results showed that the ‘traditional’ mowing regime (twice per year with the second cut during the flight period) was always detrimental to the two butterfly species at both local (single population) and regional (metapopulation) scales. However, mowing once a year, or every second or third year, before or after the flight period, was appropriate for both species in the considered landscapes. Maculinea teleius could persist only at a regional scale, assuming dispersal among the meadows, whereas M. nausithous could persist at both local and regional scales. Thus it is essential that the recommended mowing regimes are applied across several connected meadows within reach of dispersing butterflies if both butterflies are to be conserved in a region.

A Keszthelyi-hegység nappali lepkefaunája

In the course of 2003-2005 butterfly fauna of the Keszthelyi hills were studied. 1114 collected individuals belonging to 94 different species - including 23 protected species - were determined. 16 species have not been published in the butterfly fauna of the Keszthelyi hills yet. However 6 species mentioned in the literaturenot were not found during the examination. Some of the curiosities of the area are Maculinea teleius, Maculinea nausithous, Euphydryas aurinia.

Biomonitoring of the butterfly fauna in the Drava region (Lepidoptera: Diurna)

At the beginning of 2000 a biomonitoring research work started along the River Drava in Hungary. The aim of this investigation is to give information on the Hungarian biomonitoring network and up-to-date information on those environmental impacts that will also be spreading to the Drava region of Hungary, if a dam is built at Novo Virje in Croatia. A transect line estimation on the relative populations of Apatura metis, Lycaena dispar, Maculinea teleius, Maculinea nausithous, Parnassius mnemosyne, Euphydryas maturna was carried out. In this five-year period the results showed the natural fluctuation and the variability of the surveyed populations.

Microsatellite markers for the large blue butterflies Maculinea nausithous and Maculinea alcon (Lepidoptera: Lycaenidae) and their amplification in other Maculinea species

AbstractWe developed microsatellite markers for Maculinea nausithous and Maculinea alcon, two of five species of endangered large blue butterflies found in Europe. Two separate microsatellite libraries were constructed. Eleven markers were developed for M. nausithous and one for M. alcon. The primers were tested on both species as well as on the three other European Maculinea species. The number of alleles per locus ranged from two to 14. These markers will be useful tools for population genetic studies of Maculinea species.

The distribution of butterflies (Lepidoptera: Rhopalocera) in Haloze, East Slovenia

Almost nothing has been known about the butterfly fauna of Haloze prior to our survey. During our field trips, including an extensive survey carried within the framework of the Student Research Camp Videm 2002, surprisingly high butterfly diversity was established. Altogether, 96 butterfly species were recorded at 67 studied sites. The highest butterfly diversity was registered in dry grasslands in the western part of the studied area. Due to the presence of rare and endangered species, such as Maculinea arion (Linne, 1758), Meleageria daphnis (Denis & Schiffermüller, 1775), and Polyommatus thersites (Cantener, 1825), these habitats would require more conservation efforts and a suitable management. The few remaining humid grasslands in the eastern part of the region still host the majority of specialized hygrophilous butterflies, including Maculinea nausithous (Bergsträsser, 1779), Maculinea teleius (Bergsträsser, 1779), and Lycaena dispar (Haworth, 1802). Some of these sites are proposed to be included into the Natura 2000 network, which could enable their suitable protection and management.

Les Maculinea (Lepidoptera, Lycaenidae) de la Réserve naturelle du Marais de Lavours (Ain, France) et ses environs

In a previous note (Rozier, 1998) we briefly presented the natural reserve of the «marais de Lavours ». It is one of the last wetland in France where the three Maculinea'' s species : Dusky Large Blue (Maculinea nausithous ), Scarce Large Blue (M. teleius) and Alcon Blue (M. alcon) are presents. A prospecting of the xeric grassland near the reserve allowed the discovery of a fourth sort : the large blue (M. arion) making the area a zone especially rich in Maculinea.

Food-plant niche selection rather than the presence of ant nests explains oviposition patterns in the myrmecophilous butterfly genus Maculinea.

It has been suggested that the socially parasitic butterfly Maculinea alcon detects ant odours before ovipositing on initial larval food plants near colonies of its obligate ant host Myrmica ruginodis. It has also been suggested that overcrowding on food plants near M. ruginodis is avoided by an ability to detect high egg loads, resulting in a switch to selecting plants near less suitable ant species. If confirmed, this hypothesis (H1) would have serious implications for the application of current population models aimed at the conservation of endangered Maculinea species, which are based on the null hypothesis (H0) that females randomly select food plants whose flower buds are at a precise phenological stage, making oviposition independent of ants. If H1 were wrong, practical management based upon its assumptions could lead to the extinction of protected populations. We present data for the five European species of Maculinea which show that (i) each oviposits on a phenologically restricted flower-bud stage, which accounts for the apparent host-ant-mediated niche separation in sympatric populations of Maculinea nausithous and Maculinea teleius, (ii) there is no temporal shift in oviposition by Maculinea arion in relation to host ant distribution or egg density, and (iii) oviposition patterns in 13 populations of M. alcon's closest relative, Macaulinea rebeli, conform to H0 not H1 predictions. It is concluded that conservation measures should continue to be based on H0.

Comparison of genetic population structure of the large blue butterflies Maculinea nausithous and M. teleius.

We investigated the genetic population structure of two rare myrmecophilous lycaenid butterflies, Maculinea nausithous and M. teleius, which often live sympatrically and have similar biology. In Europe, both species occur in highly fragmented populations and are vulnerable to local extinction. The proportion of variable allozyme loci, average heterozygosity and genetic differentiation among populations was higher in M. nausithous than in sympatrically living M. teleius populations. We hypothesise that the differences in heterozygosity are mainly due to the known higher efficiency of typical host ant nests in rearing M. nausithous pupae compared to M. teleius pupae. This implies a larger probability of larval survival in M. nausithous, which buffers populations against environmental and demographic stochasticity. In contrast, the lower carrying capacity of ant nests in rearing M. teleius pupae requires higher nest-densities and makes M. teleius populations more prone to losing genetic variation through drift if this condition is not fulfilled. The single investigated Russian population of M. teleius showed much higher levels of heterozygosity than any of the Polish populations, suggesting a more viable and still intact metapopulation structure.

Lessons from the reintroduction of Maculinea teleius and M. nausithous in the Netherlands.

Details of a reintroduction of Maculinea teleius (scarce large blue butterfly) and Maculinea nausithous (dusky large blue butterfly) into a nature reserve in the Netherlands are given. The introduced population of M. teleius expanded during the first three years. In contrast, the newly established population of M. nausithous declined in the year after the reintroduction but expanded later. After a considerable increase in numbers, this species can nowadays be seen in two subpopulations: one in and one outside the nature reserve. Although the results are quite positive with respect to the numbers of butterflies, some alarming features have also been found. Both species, but especially M. teleius, have a very low mean minimal lifetime, which makes them more susceptible to stochastic catastrophes. While the area populated by M. nausithous is still expanding, M. teleius can only be found on the meadow where it was released and has not dispersed at all. The possible consequences of reintroductions are discussed. © Rapid Science Ltd. 1998

The behaviour and habitat requirements of Maculinea nausithous (the dusky large blue butterfly) and M. teleius (the scarce large blue) in France

Maculinea nausithous and M. teleius are among the world's rarest butterflies. Both oviposit on the flowerheads of Sanguisorba officinalis, but spend most of their immature lives inside Myrmica spp. ant nests, where their larvae feed on ant brood. Very few colonies of these butterflies survive, and leading conservation bodies have called for urgent research into their habitat requirements: at present, too little is known for successful conservation programmes to be devised. This paper described observations made on three small bogs in France, which each supported colonies of both Maculinea in 1981. The main content is a detailed habitat analysis of each site. Studies were also made of the behaviour of both butterflies, and of the size of the population of each species on every site. From these, it was possible to identify some of the factors that make these sites suitable habitats for M. nausithous and M. teleius. It eas concluded that the abundance, and perhaps the species, of Myrmica ant was the crucial factor on each site.