The effect of queen ants on the survival of Maculinea arion larvae in Myrmica ant nests

Summary The mortality of Maculinea arion caterpillars was measured in both laboratory and wild Myrmica nests, and found to be nearly 3 times higher in nests that had queen ants present. This is attributed to queen effect, which causes worker ants in nests with queens to attack large ant larvae (gynes) that would otherwise develop into new queens. Maculinea arion caterpillars mimic Myrmica larvae, and are usually attacked during the first 10 days after adoption, when they pass through the size range of ant gyne larvae. Caterpillars are also likely to be attacked during this period because their nethod of feeding brings them into close contact with the skins of large ant larvae, which contain gyne larval pheromones; older caterpillars are large enough to eat larvae without their exposed surfaces contacting the larval skin. In the wild, many caterpillars of Maculinea arion die in ant nests, and this has been shown in previous work to be the key factor that determines changes in their abundance from year to year. It is suggested that queen effect can be an important cause of these deaths, and one that particularly affects populations of butterflies that breed on sites with long-established plagioclimaxes of short turf rather than short-lived grass-land successions.     

The re-discovered Maculinea rebeli (Hirschke, 1904): Host ant usage,parasitoid and initial food plant around the type locality with taxonomical aspects (Lepidoptera,Lycaenidae)

The taxonomy of the myrmecophilous Maculinea alcon group (Lepidoptera: Lycaenidae) is highly debated. The host-plant and host-ant usage of these butterflies have conventionally been important in their identification. Maculinea ‘rebeli’ has generally been considered to be the xerophilous form of Ma. alcon (Ma. alcon X hereafter) with Gentiana cruciata as initial food plant. However, the type locality and all other known sites of Ma. rebeli are found above the coniferous zone, and are well separated from the lower regions where Ma. alcon X sites are found. Furthermore, no food plant and host ant data for the nominotypic Ma. rebeli have yet been published. Our aim was therefore to identify the host ant(s) of Ma. rebeli around the type locality and compare this with the host ant usage of nearby Ma. alcon X. Nests of Myrmica spp. (Hymenoptera: Formicidae) close to the host plants were opened on one Ma. alcon X (host plant: Gentiana cruciata) and two Ma. rebeli (host plant: Gentianella rhaetica, first record, confirmed by oviposition and emerging larvae) sites just before the flying period, to find prepupal larvae and pupae. Three Myrmica species (My. lobulicornis, My. ruginodis, My. sulcinodis) were found on the two Ma. rebeli sites, which parasitized exclusively My. sulcinodis (22 individuals in 7 nests). On the Ma. alcon X site Myrmica sabuleti and My. lonae were found, with My. sabuleti the exclusive host (51 individuals in 10 nests). Ichneumon cf. eumerus parasitized both butterflies. The results highlight the differentiation of Maculinea rebeli from Ma. alcon X, from both conservation biological and ecological points of view. Thus, it should be concluded that Ma. rebeli does not simply represent an individual form of Ma. alcon but it can be considered as at least an ecological form adapted to high mountain conditions both in its initial food plant and host ant species. In addition, it should be emphasized that Ma. alcon X (= Ma. rebeli auct. nec Hirschke) cannot be synonymised with Ma. rebeli (Hirschke, 1904).     

Population Structure, Mobility and Habitat Preferences of the Violet Copper Lycaena Helle (Lepidoptera: Lycaenidae) in Western Germany: Implications for Conservation

We describe population structure, mobility and habitat preferences of the butterfly Lycaena helle using a survey of the populations in the Westerwald area (Western Germany) and mark-recapture techniques. Three out of 83 recorded populations were intensively studied in 1995. In all, 1596 individuals (537 females) were uniquely marked. The average adult residence time was 8 days, the maximum 34 days. L. helle is extremely sedentary. Average distances between first and subsequent captures were 37 and 61 meters for males and females respectively. Population-specific differences are interpreted as being caused by the influence of barriers and corridors. The potential for colonization is regarded as poor, and isolated populations seem to be under a higher risk of extinction than other ones. Preferred habitats were abandoned moist meadows with rich aspects of Polygonum bistorta on sheltered, warm, and humid stands. Owing to the destruction of natural habitats (moorland) L. helle now colonizes almost exclusively anthropogenic ephemeral habitats (abandoned moist meadows) in Central Europe. This results in serious conservation problems, as the biology of the species does not seem to be adapted to a high turn-over rate of habitats.     

Effects of habitat reduction on the persistence of Ichneumon eumerus (Hymenoptera: Ichneumonidae), the specialist parasitoid of Maculinea rebeli (Lepidoptera: Lycaenidae)

The lycaenid butterfly, Maculinea rebeli, and its specialist parasitoid, Ichneumon eumerus, live in small, closed populations. Given the threatened status of the butterfly, it is reasonable to assume that its specialist parasitoid is even more vulnerable to local extinction than the butterfly host. Based on a mechanistic model recently developed for the tightly-woven community surrounding M. rebeli at a site in the Spanish Pyrenees, we investigate how the removal of habitat, and more particularly, specific habitat promoting the persistence of the butterfly, affects the population persistence of the parasitoid. Because of the relatively small impact of the parasitoid on the butterfly population in the Spanish Pyrenees, guidelines for conserving the parasitoid are only slightly more restrictive than those for its host. It is argued that at sites of more marginal quality for the butterfly than the reference site, achieving the dual aims of conserving both species will be more problematic. © Rapid Science Ltd. 1998     

Soldier bug predation on swallowtail caterpillars (Lepidoptera: Papilionidae): Circumvention of defensive chemistry

All larvae in the family Papilionidae possess osmeteria, eversible glands that release defensive chemicals upon disturbance. These chemicals have been shown to be repellent to ants, mantids, and other predaceous arthropods with chewing mouthparts. In this study, we demonstrate that the pentatomid Podisus maculiventris,an important predator of the black swallowtail Papilio polyxenes,is capable of piercing the body wall of the caterpillar and consuming body fluids without eliciting osmeterial eversion. Tenebrio molitorlarvae coated with osmeterial secretions of Papilio polyxeneswere universally rejected by Podisus maculiventris,suggesting that the predator is in fact repelled by the swallowtail defensive chemistry. By circumventing the defensive behavior of the caterpillar, the pentatomid is able to utilize an otherwise chemically unsuitable prey species.     

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

We 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.     

Non-resource based territoriality in males of the butterflyXamia xami (Lepidoptera: Lycaenidae)

In the Pedregal de San Angel reserve, in Mexico City, males of the butterfly Xamia xamiperch in and defend areas with well-defined topographic limits. These areas lack concentrations of receptive females and of larvae and adult resources. One individual defends the same territory an average of 5 h/day, up to a maximum of 23 days. The same areas are used as territories by different males during the year. These areas share some characteristic features which are described. Evidence is presented in support of the hypothesis that the territories function as mating stations. A possible scenario for the evolution of this territorial mating system is advanced.     

Acid stress in Rumex hydrolapathum (Polygonaceae) and its influence on the phytophage Lycaena dispar (Lepidoptera; Lycaenidae)

Summary The relationship of insect growth to host condition was tested on host plants grown by hydroculture with a standard nutrient solution, but under different acidities, ranging from pH 3.5–7.5. The indirect effect of the host on the development of the phytophage was tested in the pupal stage. The host plant Rumex hydrolapathum had its highest nitrogen content in the range pH 5.5–6.5; pupal weight of Lycaena dispar was correlated with nitrogen concentration, ash weight and water content. There is a remarkable difference in element concentration between young and old leaves of the host, which varies with acidity of the rooting substrate.In this experiment a negative relationship was detected between reproduction of the phytophage and stress of the host.     

Experimental Extinction of Garlic Mustard (Alliaria petiolata) Populations: Implications for Weed Science and Conservation Biology

An assumption of weed science and conservation biology is that small populations are more vulnerable to elimination and extinction than large populations. We tested this with the invasive biennial garlic mustard (Alliaria petiolata). We compared 61 experimental populations from which every flowering plant was removed for 4 years, with 56 control populations. Whereas the majority of the control populations continued to expand in size over the 4 years, experimental populations showed a strong experimental effect, remaining stable in size, declining in size, or going extinct. Small populations were far more vulnerable to extinction than large populations: 43% of small experimental populations (initially fewer than 10 individuals) went extinct, but only 7% of large populations (initially more than 50 individuals). However, some small experimental populations persisted, and in a few cases, larger experimental populations continued to expand even though every flowering individual had been removed. These results and a simple population model suggest the importance of buried seeds in allowing this species to persist despite attempts to eradicate it.     

Multilevel studies on the two phenological forms of Large Blue (Maculinea arion) (Lepidoptera: Lycaenidae)

The main goal of our research was to study comprehensively the differences between the two phenological forms of the socially parasitic and globally threatened Large Blue (Maculinea arion) in the Carpathian Basin using four character sets (mitochondrial sequences, allozymes, male genitalia and wing morphometrics). Comparative analyses of distance matrices, phylogenetic trees and ordination patterns have been applied. The genetic and morphometric patterns revealed by our studies were discordant. While we experienced a significant differentiation between the ‘spring’ and ‘summer type’ of M. arion in both wing and genital traits, the two phenological forms did not show any genetic differentiation on two mitochondrial loci and in allozymes. At the same time, all individuals were infected by Wolbachia. Although certain wing traits may not represent reliable tracers of phylogeny because of the particular adaptive significance, the wing characteristics involved in our research are probably determined genetically. Additionally, the significant differentiation of male genitalia also indicates incipient prezygotic isolation arising from phenological differentiation between the ‘spring and summer arion’. It is possible that all extant differences between the two forms are attributable to (1) different host‐ant use, (2) incipient speciation, (3) cytoplasmatic incompatibility (CI) by Wolbachia or the combination of these factors. In addition, discordant results indicate that the combined use of different approaches and data sets is strictly necessary to clarify systematic and evolutionary relationships.     

Reconstructing intraordinal relationships in Lepidoptera using mitochondrial genome data with the description of two newly sequenced lycaenids, Spindasis takanonis and Protantigius superans (Lepidoptera: Lycaenidae)

Lepidoptera is one of the largest insect orders, but the phylogenetic relationships within this order, have yet to be adequately described. Among these unresolved relationships include those regarding the monophyly of the Macrolepidoptera and interfamilial relationships of the true butterflies superfamily Papilionoidea. We present two new mitochondrial genomes (mitogenomes) belonging to the butterfly family Lycaenidae to explore the phylogenetic relationships existing among lepidopteran superfamilies and true butterfly families from a mitogenome perspective, and to evaluate the characteristics of the lepidopteran mitogenomes. Our consensus phylogeny of the Lepidoptera largely supported the superfamilial relationships (((((Bombycoidea + Geometroidea) + Noctuoidea) + Pyraloidea) + Papilionoidea) + Tortricoidea), signifying a lack of support for a traditionally defined Macrolepidoptera. The familial relationships of the true butterflies concordantly recovered the previously proposed phylogenetic hypothesis (((Lycaenidae + Nymphalidae) + Pieridae) + Papilionidae). The test for the effect of optimization schemes (exclusion and inclusion of third codon position of PCGs and two rRNA genes, with and without partitions) on the resolution and relationships within the Lepidoptera have demonstrated that the majority of analyses did not substantially alter the relevant topology and node support, possibly as the result of relatively strong signal in mitogenomes for intraordinal relationships in Lepidoptera.     

Somatic embryogenesis of <Emphasis Type="Italic">Gentiana cruciata</Emphasis> (L.): Histological and ultrastructural changes in seedling hypocotyl explant

Summary Structure and ultrastructure changes that occurred during tissue culture of upper explants of hypocotyl (adjacent to cotyledons) of 10-d-old seedlings of Gentiana cruciata were studied. The explants were cultured on Murashige and Skoog induction medium supplemented with 1.0 mg l⁻¹ dicamba +0.1 mg l⁻¹ naphthaleneacetic acid +2.0 mg l⁻¹ benzyladenine +80.0 mg l⁻¹ adenine sulfate. The initial response of the explant and callus formation were ultrastructurally analyzed during the first 11 d of culture. After 6–8 wk, various methods were employed to collect evidence of indirect somatic embryogenesis. After 48 h of culture, the earliest cell response was cell division of epidermis and primary cortex. There were numerous disturbances of karyo- and cytokinesis, leading to formation of multinuclear cells. With time, the divisions ceased, and cortex cells underwent strong expansion, vacuolization and degradation. About the 6th day of culture, callus tissue proliferated and the initial divisions of vascular cylinder cells were observed. Their division appeared normal. Cells originating from that tissue were small, weakly vacuolated, with dense cytoplasm containing active-looking cell organelles. Numerous divisions occurred in the vascular cylinder, which led to its expansion and the formation of embryogenic callus tissue. During the 6–8th wk of culture, in the proximal end of the explant, masses of somatic embryos were formed from outer parts of intensively proliferating tissue.     

Susceptibility of the ant-cultivated fungus Leucoagaricus gongylophorus (Agaricales: Basidiomycota) towards microfungi

The aim of this study was to select virulent strains of microfungi against Leucoagaricus gongylophorus, a symbiotic fungus cultivated by leaf-cutting ants. The results from in vitro assays showed that microfungal strains had a variable and significant impact on the colony development of L. gongylophorus. Specifically, Trichoderma harzianum, Escovopsis weberi CBS 810.71 and E. weberi A088 were more effective, inhibiting the L. gongylophorus colonies by 75, 68 and 67%, respectively (P < 0.05) after 15 days. Strain E. weberi A086 and Acremonium kiliense were less effective: 43 and 26%, respectively (P < 0.05). In spite of the current negative perspective of a microbiological control approach for these ants, the present work discusses the possibility of using mycopathogenic fungi for the control of these insects, and points out the importance of encouraging more studies in this area.     

The influence of diet on growth and secretion behaviour of myrmecophilous Polyommatus icarus caterpillars (Lepidoptera: Lycaenidae)

Abstract.

• 1 Caterpillars of the facultatively myrmecophilous butterfly Polyommatus icarus were reared on inflorescences, or foliage, of four natural hostplant species and on an artificial diet to study dietary effects on larval growth and secretory capacity.
• 2 Caterpillars achieved highest weights and relative growth rates when fed flowers of Medicago sativa, Lotus corniculatus or Melilotus officinalis. Larvae reared on Coroniüa varia (flowers and leaves), foliage of M.sativa and on the artificial diet pupated at lower weights and achieved lower growth rates.
• 3 In standardized experiments with the ant species Lasius flavus, secretion rates from the dorsal nectar organ (DNO) were 2 times higher among flower-fed caterpillars than among foliage-fed siblings or caterpillars on the artificial diet. Larvae reared on C.varia flowers were superior to all other food treatments with respect to secretion rates.
• 4 High water content of larval diet, as in flowers, appears to be important for lycaenid caterpillars to achieve high secretion rates, whereas the correlation between myrmecophily and nutrient availability, as evidenced by growth rates, was less pronounced.
• 5 Using experimental data on larval growth and secretion rates, the lifetime volume of secretions from the DNO is estimated to range from 2 to 5 μl in most food treatments. Only on C.varia flowers (5.5–8.7 μl) and on M.sativa leaves (0.9–1.1 μl) did the caterpillars deviate in their absolute investment in myrmecophily.
• 6 The estimated lifetime investment accounted for 1.6–5.5% of prepupal fresh weight in all food treatments except on C.varia flowers (7.8–12.3%).

     

The fragmented population structure of the Sardinian chalk hill blue butterfly (Lepidoptera, Lycaenidae)

The endemic Sardinian chalk hill blue butterfly, Polyommatus coridon gennargenti, is considered vulnerable to extinction because of its low genetic variation and restricted distribution. The species also has a fragmented distribution, which follows the patchy distribution pattern of its larval host-plant. A preliminary investigation of the population structure of P. coridon gennargenti was carried out on a small network of four local populations by means of capture–recapture methods. Estimated population sizes and movement rates among the four adjacent local populations suggest that this taxon has a metapopulation structure composed of loosely connected small local populations. Natural fragmentation, isolation, and traditional land use contribute to the vulnerability of P. coridon gennargenti to extinction. Low effective population sizes and restricted movement between habitat patches lead to inbreeding and an increased vulnerability to extinction of this island population.     

Characterization of microsatellite loci in the black rhinoceros (Diceros bicornis) and white rhinoceros (Ceratotherium simum): their use for cross-species amplification and differentiation between the two species

As the population sizes of the black and white rhinoceroses continues to decline, more efforts are needed in multiple areas to help with the conservation efforts. One area being explored is the use of genetic diversity information to aid conservation decisions. In this study, we designed 21 microsatellite primers for white and black rhinoceroses, 16 and 17 of which amplified bands in the white and black rhinoceros, respectively. Out of these primers all 16 were polymorphic in the white rhinoceros and 12 of the 17 were polymorphic in the black rhinoceros. The mean number of alleles was 3.31 and 2.12, the expected heterozygosities were 0.420 and 0.372, and the observed heterozygosities were 0.436 and 0.322 for the white and black rhinoceroses, respectively. Seven of the primers produced different allele sizes and variations that distinguished between black and white rhinoceroses. Further genetic analyses with larger wild population sample sizes and markers are recommended to obtain a better understanding of the genetic structure of the black and white rhinoceros populations in order to be useful in the conservation efforts of these critically endangered species. A. Kilbourn—In memoriam.     

Conservation management of Gortyna borelii lunata (Lepidoptera: Noctuidae) in the United Kingdom

Gortyna borelii lunata is a rare species of moth with a widespread, but localised distribution in Europe. In Britain, the moth is restricted to coastal grassland habitats that support its larval foodplant (Peucedanum officinale), in the southeast of England.Threats to G. borelii lunata in Britain include inappropriate management and sea-level rise. This study investigates how to secure the future of the moth in Britain by managing existing colonies appropriately and determining whether translocations of the species, away from the dangers of flooding, would be feasible. A mowing experiment and a translocation trial were conducted. Results from the mowing experiment indicate that cutting annually in either August or November is detrimental to the abundance of the moth. It is recommended that where sites must be mown this be performed on rotation, leaving much of the site uncut each year. The translocation of the moth and its foodplant to a site away from the threats of flooding was very successful, demonstrating that the species establishes well and can survive on higher ground. It is concluded that to secure the future of the moth in Britain, a sustainable network of interconnected sites for the species should be created. Also, an appropriate management plan should be formulated for each site.     

Comparing the effects of the exotic cactus-feeding moth, Cactoblastis cactorum (Berg) (Lepidoptera: Pyralidae) and the native cactus-feeding moth, Melitara prodenialis (Walker) (Lepidoptera: Pyralidae) on two species of Florida Opuntia

This study examined the effects of the native cactus moth borer, Melitara prodenialis, and the invasive cactus moth borer, Cactoblastis cactorum, on two common cactus species, Opuntia stricta and O. humifusa at coastal and inland locations in central Florida. Opuntia stricta were present only at coastal sites and O. humifusa were present at coastal and inland sites. Throughout the duration of the study, coastal plants were subject to damage solely by C. cactorum and inland plants solely by M. prodenialis. Results showed marginally significantly higher numbers of eggsticks on O. stricta than O. humifusa and significantly higher numbers at coastal sites than at inland sites. There was also significantly higher moth damage on O. stricta than O. humifusa and at coastal sites than inland sites, but not significantly so. However, there was a higher level of plant mortality for O. humifusa than for O. stricta and a significantly higher level of cactus mortality at inland sites when compared to coastal sites. This increased mortality may be due to increased attack by true bugs, Chelinidea vittiger, and by Dactylopius sp., combined with attack by M. prodenialis. Inland plants also tended to be smaller than coastal plants and could be more susceptible to the combined effects of all insects. Further long-term research on coastal cactus survival when attacked and unattacked by Cactoblastis is necessary to fully determine the effects of this moth on Opuntia survival.     

温度对荔枝异形小卷蛾发育和繁殖的影响

荔枝异形小卷蛾的蛀梢为害是珍贵树种格木人工林健康发展的主要限制因子。依据其天然分布和潜在推广区的温度范围设置系列温度梯度,探讨温度对荔枝异形小卷蛾发育和繁殖的综合影响。结果显示,温度对荔枝异形小卷蛾各阶段的发育历期具有显著影响,在研究温度范围内,发育历期随温度升高呈显著地下降趋势,世代历期在18℃时为66.87 d,30℃时降至35.77 d。预蛹和蛹的存活率对温度的响应不敏感,而卵、幼虫、成虫的存活率和世代存活率以及产卵量均随温度升高表现为先上升后下降的变化趋势,且繁殖力对温度的反映较存活率敏感,其存活率和繁殖力在18℃时均最低,分别为41.20%和13.90粒,在27℃时发育最适,分别为83.80%和45.40粒,在30℃时虽有下降,分别为66.00%和32.40粒,但仍高于18℃时,即其对低温较高温敏感。荔枝异形小卷蛾完整世代发育起点温度为5.77℃,所需有效积温为876.76 d℃,其中幼虫发育所需有效积温最高,占整个历期的45.23%,发育速率与温度显著正相关。根据荔枝异形小卷蛾为害方式、发育和繁殖特征分析可知,在低温地区其幼虫期长,但世代数少、存活率和繁殖力低,对寄主植物受害部位的单次为害程度严重;在高温地区则幼虫期短,但世代数多、存活率和繁殖力高,对寄主植物受害部位的单次为害程度低,但更为频繁,持续为害程度高。研究结果对于不同地区选择荔枝异形小卷蛾的防治具有借鉴意义,同时有助于指导格木人工林的合理推广和健康发展。