Biology and conservation of <Emphasis Type="Italic">Zerynthia cassandra</Emphasis> (Lepidoptera,Papilionidae) in semi-natural environments and agricultural landscapes

The Southern festoon, Zerynthia polyxena, is a south-European butterfly listed in Annex IV of the Habitats Directive (92/43/EEC). Populations from most of the Italian territory have been recently recognized as belonging to a separate species, endemic to Italy: Zerynthia cassandra. In the province of Bologna (northern Italy), Z. cassandra is quite common. The territory of the province is densely inhabited and modified by man. Semi-natural environments are found almost exclusively in the hills, while the plain is almost entirely built-upon or cultivated. This study was based on data collected in the field in the province of Bologna, and aimed to investigate some biological and ecological aspects of Z. cassandra, with the purpose of identifying the factors that favour its presence and the most relevant issues for its conservation. In the area of study, the hostplant of Z. cassandra is Aristolochia rotunda, which is common in the plain, along the edges of drainage canals, where it supports a large Z. cassandra population. In the hills, A. rotunda is scarcer, and Z. cassandra is present in small separated nodes. Human activities such as mowing and canal maintenance are potentially damaging to Z. cassandra, particularly if carried out when eggs and larvae are present. However, if correctly timed and executed, such maintenance can preserve grassy areas that would otherwise be colonized by bushes and trees, and in which A. rotunda finds a suitable environment. Some level of disturbance can be beneficial for Z. cassandra habitats, favouring the species’ survival in the long term.     

Mobility and oviposition site-selection in Zerynthia cassandra (Lepidoptera,Papilionidae): implications for its conservation

The adults’ mobility and oviposition preferences of Zerynthia cassandra have been studied for the first time, with the aim of integrating auto-ecological information into recommendations for the habitat’s management of this species. Results of our mark-release-recapture study have highlighted that Z. cassandra is a strictly sedentary species, since detected movements only occurred over very short distances (≤200 m) and mainly within the species’ reproductive habitat (i.e. around Aristolochia rotunda stands), with males moving further than females. Our study shows that the main oviposition habitat of Z. cassandra is found where A. rotunda plants are growing in large stands; sites where plants growing in half to full sun and mostly oriented to the south are preferred. The distance of deposited eggs from the plants’ roots was narrowly correlated with the plants’ length. Eggs were deposited singly, mainly on the underside of leaflets. Management strategies necessary for improving the most important habitat features for the conservation of this species are suggested.     

Consequences of the arms race between <Emphasis Type="Italic">Maculinea teleius</Emphasis> social parasite and <Emphasis Type="Italic">Myrmica</Emphasis> host ants for myrmecophilous butterfly conservation

The arms race between Maculinea butterflies and Myrmica host ants leads to local host-parasite adaptations. In our study, we assessed whether sympatric and allopatric Myrmica scabrinodis populations exhibit behavioural differences towards Maculinea teleius larvae during the adoption-period when butterfly larvae need to be taken inside the Myrmica nest. The second aim was to assess the butterfly survival rate inside ant colonies from different populations. We used one sympatric host population and three allopatric populations: one infested by M. teleius and two uninfested populations. We found that ants from the sympatric population showed a higher number of positive behaviours toward M. teleius larvae during adoption than ants from the allopatric populations. There were no differences in the number of inspection or negative behaviour events. The survival of butterfly larvae was highest inside sympatric host colonies and differed from the survival of M. teleius reared by ants from the allopatric, uninfested populations. No difference was found for the survival rate of M. teleius raised by infested, allopatric host colonies compared to sympatric host populations. Our results suggest the lack of behavioural counter-adaptations of local hosts of M. teleius that more easily adopt and rear butterfly caterpillars compared to naive M. scabrinodis colonies. Our results may also have implications for Maculinea butterfly conservation, especially for reintroduction programmes. We suggest that the existence of behavioural host defences should be checked for the source host population, as well as for the Myrmica population from the reintroduction site. It may also be reasonable to introduce several Myrmica host colonies from the source butterfly host population.     

Fen meadows on the move for the conservation of <Emphasis Type="Italic">Maculinea (Phengaris) teleius</Emphasis> butterflies

In the Netherlands, a single population of the obligate myrmecophilic butterfly Maculinea (Phengaris) teleius has survived on only 3 ha of habitat for more than 25 years, whereas at least 40 ha of habitat are thought to be required for a sustainable metapopulation. Therefore, 170 ha of farmland is being restored to wet meadows within a LIFE?+?project by large-scale soil excavation and hay inoculation. For successful restoration, the habitat requirements of the butterfly, with Sanguisorba officinalis as host plant and its particular life cycle as parasite of the ant species Myrmica scabrinodis, have to be taken into account. We tested whether colonization of nests of this ant species in the restoration areas is facilitated by translocation of sods collected from fen meadows. We divided 54 sods, each sized 1 m², randomly over six patches and measured vegetation development and ant presence in the sods and surrounding control plots for 2 years. In the first summer, significantly more Myrmica ants were found in the transplanted sods in comparison to the surrounding area. Herb cover had a significant positive effect on Myrmica ant presence while it did not affect the presence of the pioneer ant species Lasius niger. In the second year, Myrmica ants were found in the surrounding control plots as well. This study contributes to the knowledge-base required for the design of restoration projects aimed at expanding the habitat of the critically endangered butterfly Maculinea (Phengaris) teleius.     

Oviposition site selection of the Alcon blue butterfly at the northern range margin

Female oviposition decisions in insects may strongly affect offspring growth and survival, and thus determine population performance. In this study, we examined oviposition site selection in the xerophilous ecotype of the endangered myrmecophilous butterfly Phengaris (=Maculinea) alcon (Lepidoptera, Lycaenidae) in Estonia, at the northern distribution margin of both the butterfly and its host plant, Gentiana cruciata L. Egg distribution on individual host plants appeared to be highly uneven: plants carrying high egg loads contrasted to a high proportion of host plants without any or with a few eggs. Host plant use for oviposition was strongly dependent on plant characteristics and environmental context. Host plant height relative to the surrounding vegetation rather than the absolute height of host plants was a key factor determining the use of particular host plant individuals for oviposition. In particular, plants protruding above surrounding vegetation had a higher probability of being used for oviposition, and carried more eggs. Additionally, the number of eggs laid on individual host plants was positively associated with the presence of flowers and the number of shoots. More aggregated host plants received fewer eggs than those with less conspecifics around. Feeding damage by wild herbivores, found in a substantial proportion of the butterfly’s host plants, strongly reduced the number of eggs on individual plants. Our results underline the need to assure that butterfly’s host plants do not become overgrown by surrounding vegetation. Best practices for opening vegetation around host plants may need further studies that explicitly account for butterfly’s host ants—their abundance in relation to vegetation height and their response to opening vegetation.     

Is It Possible to Manipulate Scelionidae Wasps’ Preference to a Target Host?

Parasitoid host selection is mainly mediated by chemical cues, which can be adjusted by experience, changing their innate behavior. Therefore, this study evaluated if immature experience (pre-imaginal conditioning) on eggs and volatiles from different host eggs has influence on parasitism and chemotaxic behavior of Telenomus podisi Ashmead and/or Trissolcus basalis Wollaston (Hymenoptera: Scelionidae). Both wasp species were submitted to a multiple-choice parasitism test among Euschistus heros (Fabricius), Piezodorus guildinii (Westwood), and Nezara viridula L. (Hemiptera: Pentatomidae) (Hemiptera: Pentatomidae) egg masses. Eggs from these three stink bugs were equally offered to female parasitoids. After that, adults which emerged from each host were also exposed to parasitism in a multiple-choice test for up to an additional generation. Moreover, in olfactometer “Y,” the behavior of innate and experienced T. podisi females to volatiles from hosts’ egg extracts was tested, to study their learning and memory ability. The original host had influence on T. podisi parasitism; however, T. basalis always parasitized more N. viridula eggs independently of its last rearing host. Innate T. podisi females responded positively to E. heros and P. guildinii egg volatiles, but this behavior was not observed in N. viridula. When T. podisi females were experienced on egg volatiles from a new host, they showed significant learning and memory ability for the specific host volatile for, at least, 24 h. Experienced wasps responded positively to N. viridula and through this result we have evidences about the possibility to manipulate wasp’s preferences to a specific target host.     

Effect of experience on response of <Emphasis Type="Italic">Trichogramma buesi</Emphasis> Voeg. and <Emphasis Type="Italic">T. principium</Emphasis> Sug. et Sor. (Hymenoptera,Trichogrammatidae) females to different ages of host eggs

Laboratory experiments have demonstrated that patterns of distribution of T. principium and T. buesi females by the number of mature ovarial eggs is strongly dependent on the age of the host (Sitotroga cerealella) eggs offered for parasitization. The approximate percentage of parasitizing females estimated from these distributions was 1.5–2 times higher for females offered young host eggs than for those offered “old” eggs, that had developed for six days at a temperature of 20°C. In both species, females more often oviposited in old (non-preferred) eggs when they had previously experienced fresh (preferred) eggs. This result contradicts the widely accepted assumption that in insect parasitoids, experience of parasitizing a preferred host decreases the probability of parasitizing a non-preferred host. One might expect that under natural conditions, a rather long time interval also precedes parasitization of less preferred host eggs by Trichogramma females, and duration of this interval may be also dependent on availability of other host species.     

Rate of consumption,biological parameters,and population growth capacity of <Emphasis Type="Italic">Orius laevigatus</Emphasis> fed on <Emphasis Type="Italic">Spodoptera exigua</Emphasis>

Orius laevigatus (Fieber) (Hemiptera: Anthocoridae) is already successfully used to control Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae) in protected crops. In the present work, the predatory capacity of O. laevigatus on the eggs and larvae of Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) as well as the population growth capacity of O. laevigatus fed on eggs of the noctuid were determined. Fourth and 5th instar nymphs had a greater consumption of eggs than younger ones. Females consumed more eggs than males. Only 5th instar nymphs and adults preyed on the neonatal larvae of S. exigua. The intrinsic rate of natural increase did not differ between O. laevigatus fed with S. exigua eggs and those offered eggs of the substitute host Ephestia kuehniella (Zeller) (Lepidoptera: Pyralidae). Thus, O. laevigatus is a good candidate for the biological control of S. exigua, a cosmopolitan pest of many crops.     

Patch occupancy and habitat of the hops azure (<Emphasis Type="Italic">Celastrina</Emphasis><Emphasis Type="Italic">humulus</Emphasis>), a rare North American endemic butterfly: insights for monitoring and conservation

The hops azure (Celastrina humulus Scott & D. Wright 1998) is a rare butterfly found along the Front Range of Colorado. Data on the prevalence of the butterfly and its preferred habitats are lacking. To describe the habitat of C. humulus at the southern part of its range, explore what factors impact C. humulus detectability, and estimate C. humulus habitat use along a riparian area known to support it, we conducted an occupancy analysis along the largest riparian system at the U.S. Air Force Academy (USAFA) in Colorado, USA. We used environmental and site-specific covariates to model the probability of detection and the probability of occupancy. Probability of detecting C. humulus was influenced by the amount of cloud cover during sampling, while the probability of occupancy was influenced by the total area of the host plant (wild hops) at the site. Probability of detection was higher during the first visit (69 %) than the second visit (64 %), and the probability of occupancy was higher (77 %) than assumed (30 %). Despite the host plant being patchily distributed throughout the butterfly’s range, the riparian areas at USAFA had a high prevalence of both wild hops and C. humulus. We use the project findings to develop future sampling efforts for the butterfly along tributaries at other locales within the butterfly’s range.     

Host suitability of the Mediterranean flour moth for rearing <Emphasis Type="Italic">Meteorus pulchricornis</Emphasis> (Hymenoptera: Braconidae), a polyphagous endoparasitoid of pest lepidopteran larvae

We investigated the host suitability of the Mediterranean flour moth Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) for a polyphagous koinobiont endoparasitoid Meteorus pulchricornis (Wesmael) (Hymenoptera: Braconidae), a common natural enemy of various pest lepidopteran larvae. The estimated probability of adult wasp emergence was 80% or higher when eggs were laid in nearly fully grown larvae of E. kuehniella (fresh weight, >?20.0 mg). The body size of emerged adult wasps increased with the initial weight of the host larvae at oviposition. The fresh weight of adult wasps reared on E. kuehniella was approximately 60% of that when reared on a natural host Spodoptera litura (Fabricius) (Lepidoptera: Noctuidae), and the lifetime fecundity of wasps reared on E. kuehniella was approximately half of that when reared on S. litura. Ephestia kuehniella was shown to be a positive host candidate for the mass rearing of M. pulchricornis, but further investigation is needed to increase the body size of wasps for more practical use of this species as a biocontrol agent.     

Reproductive Behavior of the Bug <Emphasis Type="Italic">Molipteryx fuliginosa</Emphasis> Uhler (Heteroptera,Coreidae) in the South of the Russian Far East

Reproductive behavior of Molipteryx fuliginosa (Uhler) was investigated in Primorskii Territory of Russia. From 4 to 18 repeated copulations of one female lasting from 2 to 48 hours were recorded in cages. The behavior of ovipositing females and the stages of oviposition are described for the first time. The number of eggs laid between copulations varied from 1 to 13, the number of oviposition acts, from 4 to 11, and the total female fecundity, from 21 to 38 eggs. Caged females laid eggs on plants and also on dead substrates unsuitable for nymphal feeding, such as cloth, dry branches, and a wooden pole. Copulation of M. fuliginosa was also observed under natural conditions. The preferred mating places of M. fuliginosa in anthropogenically modified habitats and in small-leaved riparian forests were plants of Rubus idaeus L., R. caesius L., and Rubus sp. After mating, females migrated in search of places for oviposition. Single eggs were found on the following plants not known previously as hosts of this bug: Solanum lycopersicum L., Carex sp., Elytrigia repens (L.) Nevski, and Taraxacum officinale Wigg. The females seemed to lack selectivity in the choice of place for oviposition, which was not always associated with host plants, despite their abundance and availability.     

How do two specialist butterflies determine growth and biomass of a shared host plant?

Although insect herbivory can modify subsequent quantity and quality of their host plants, change in plant quantity following herbivory has received less attention than plant quality. In particular, little is known about how previous herbivore damage determines plant growth and biomass in an insect species-specific manner. We explored whether herbivore species-specific food demand influences plant growth and biomass. To do this, we conducted a series of experiments and field survey using two specialist butterflies, Sericinus montela and Atrophaneura alcinous, and their host plant, Aristolochia debilis. It is known that A. alcinous larva requires four times more food resources to fulfill its development than S. montela larva. Despite that A. alcinous larvae imposed greater damage on plants than S. montela larvae, plant growth did not differ due to herbivory by these species both in single and multiple herbivory events. On the other hand, total aboveground biomass of the plants was reduced more by A. alcinous than S. montela feeding regardless of the number of herbivory events. Feeding on plants with a history of previous herbivory neither decreased nor increased larval growth. Our results suggest that food demand of the two butterfly species determined subsequent plant biomass, although the plant response may depend on tolerance of the host plant (i.e., ability to compensate for herbivore damage). Such difference in the effects of different herbivore species on host plant biomass is more likely to occur than previously thought, because food demand differs in most herbivore species sharing a host plant.     

Environmental risk assessment of the egg parasitoid <Emphasis Type="Italic">Anaphes inexpectatus</Emphasis> for classical biological control of the <Emphasis Type="Italic">Eucalyptus</Emphasis> snout beetle, <Emphasis Type="Italic">Gonipterus platensis</Emphasis>

Classical biological control is a valuable tool against invasive pests, but concerns about non-target effects requires risk assessment studies. Potential non-target effects of Anaphes inexpectatus Huber and Prinsloo (Hymenoptera: Mymaridae) were assessed for a classical biological control programme against the Eucalyptus snout beetle, Gonipterus platensis (Marelli) (Coleoptera: Curculionidae). No-choice tests were conducted with 17 non-target species to assess host specificity, including 11 curculionids. In behavioural observations, A. inexpectatus showed no interest in any of the non-target species, but two weevil species were parasitised within five days of exposure, although at significantly lower rates than G. platensis. In choice tests, only one non-target, Hypera postica (Gyllenhal) (Coleoptera: Curculionidae), was parasitised, at a rate of 0.6%, while 50.0% of G. platensis eggs were parasitised. Based on the host specificity test results and the potential host fauna found in the target area, the likelihood of non-target effects resulting from the release of A. inexpectatus is considered to be negligible.     

Host-plant patch qualities and presence of a likely competitor species affect the distribution and abundance of a rare British moth, <Emphasis Type="Italic">Cucullia lychnitis</Emphasis>

Understanding the environmental factors that affect the distribution and abundance of rare and threatened species can help identify priority sites for conservation action such as habitat management. The moth Cucullia lychnitis is identified in UK legislation as a priority species for conservation. Its conspicuous larvae feed on a patchily distributed host plant Verbascum nigrum, but are entirely absent from some host plant patches within the species’ range. Environmental variability among patches was investigated to understand factors affecting (i) patch occupancy by C. lychnitis and (ii) abundance of C. lychnitis, where present. Occupancy of individual V. nigrum plants within occupied patches was also investigated. The likelihood of patch occupancy increased with patch size, and decreased with patch isolation. Abundance of C. lychnitis was negatively correlated with patch isolation and with abundance of the weevil Cionus nigritarsis, which exploits the same host plant and may be a competitor. Within occupied patches, larvae of C. lychnitis were significantly more likely to be found on taller plants with more flower spikes; such plants are typical of established rather than newly-created patches. These results may help to guide efforts to conserve C. lychnitis, highlighting the importance of habitat connectivity and indicating potential management actions to promote features positively associated with larval incidence and abundance.     

Patterns of microhabitat and larval host-plant use by an imperiled butterfly in northern Florida

The quality of habitat for a given species is fundamental to its persistence in that habitat space. Herbivorous insects often require a specific combination of host plants, floral resources, and physical features such as shelter. Identifying these different habitat features is a focus of ecology and conservation, particularly for managing rare or imperiled taxa. We investigated the patterns of microhabitat and host plant use of the rare frosted elfin butterfly, Callophrys irus, a larval host-plant specialist found in frequently disturbed sand plains, barrens, and sandhill pine-oak forests of the eastern United States. Previous studies have been conducted on populations in the Northeastern and Midwestern US, but the southern part of its range remains unstudied. Our efforts focused on a persistent C. irus colony in northeastern Florida, resulting in a geographically referenced census of larval host-plant Lupinus perennis, along with a multiple year survey of microhabitat features relevant to both C. irus adults and immatures. Results of the larval host-plant census revealed that the highest densities of host plants were located distant to the highest densities of C. irus. Hot-spot analysis confirmed the significance of this pattern, suggesting different habitat requirements for larval host-plant L. perennis and C. irus individuals in order to achieve maximum potential densities. Our survey of C. irus immatures showed a similar pattern of microhabitat affinity that was previously recorded in the Northeast and Midwestern US, including large larval host-plants, low amounts of ground cover vegetation, and the presence of some shade. Unique to our study we found that the presence of other herbivores of L. perennis such as larvae of the crambid moth Uresiphita reversalis had a negative effect on encountering C. irus immatures. Our results suggest that management that aims to conserve these species needs to include habitat factors that favor the overlap of these species and to consider where their densities are the highest.     

Effects of Host Interspecific Interaction in the <Emphasis Type="Italic">Maculinea</Emphasis>–<Emphasis Type="Italic">Myrmica</Emphasis> Parasite–Host System

A model of interspecific host competition in a system with one parasite (butterfly—Maculinea) and multiple potential hosts (ants—Myrmica) is presented. Results indicate that host interspecific competition increases the occurrence of multiple host behaviour in Maculinea natural populations but decreases the ability of the parasite populations to adapt to the most abundant host species. These qualitative predictions were compared with data on host specificity, with good agreement. Analysis of the data also indicates that Maculinea teleius and Maculinea arion respond differently to changes in relative host abundances. Maculinea teleius shows a larger fraction of sites where it displays multiple host behaviour and a larger fraction of sites where the niches of the hosts overlap. In some instances, Maculinea teleius is adapted to Myrmica hosts that are present in lower frequencies. Maculinea arion is locally more host-specific and occurs at sites where host interspecific competition is unlikely and is more frequently adapted to the most abundant host species.     

Chemical defences of native European coccinellid eggs against intraguild predation by the invasive Asian coccinellid, <Emphasis Type="Italic">Harmonia axyridis</Emphasis> (Pallas)

Harmonia axyridis (Pallas) is a coccinellid of Asian origin that has recently invaded substantial parts of Europe and is suspected to affect native coccinellid populations through intraguild predation and competition for food. Previous work has shown that two species from the Calvia genus appeared to be well protected against H. axyridis predation. To deepen our understanding on chemical protection of Calvia spp. and the predation risk by H. axyridis, we tested for susceptibility and palatability of Calvia spp. and H. axyridis eggs against predation by H. axyridis neonate larvae. Results show that eggs of C. quatuordecimguttata were mostly not eaten by H. axyridis, while eggs of the congeneric C. decemguttata were found to be largely unprotected against predation by the invasive coccinellid. We also observed that H. axyridis first instars successfully cannibalized on conspecific eggs. Removing the surface chemicals from C. quatuordecimguttata eggs resulted in significantly reduced protection from being preyed upon by H. axyridis, while applying these extracts onto C. decemguttata and H. axyridis eggs resulted in increased protection against H. axyridis larvae. The importance of surface chemicals in the interactions between H. axyridis and native coccinellids was confirmed by GC–MS analysis, showing a high diversity of hydrocarbons located on the surface of C. quatuordecimguttata eggs, i.e. more than twice as many when compared to C. decemguttata. Survival of H. axyridis larvae feeding on eggs of C. quatuordecimguttata, C. decemguttata or conspecific eggs, from which surface chemicals were removed by washing them with hexane, was not different from survival on unwashed eggs.     

Allozyme variation of the relict plant <Emphasis Type="Italic">Aristolochia manshuriensis</Emphasis> Kom. (Aristolochiaceae)

Allozyme variation of a rare relict plant, birthwort Aristolochia manshuriensis Kom., was examined. The main parameters of genetic variation in natural populations of A. manshuriensis from the Anan’evka, Nezhinka, and Malaya Borisovka river basins (Primorskii Krai) were inferred from analysis of nine enzyme systems, presumably encoded by 18 loci. At 99% polymorphism criterion, 24.4% of A. manshuriensis loci were shown to be polymorphic. The mean number of alleles per locus was 1.24; the mean observed and expected heterozygosities, 0.12 and 0.10, respectively.     

An endangered hemiparasitic shrub is the only host plant of the little-known Neotropical hairstreak <Emphasis Type="Italic">Strymon flavaria</Emphasis> (Lepidoptera: Lycaenidae) in the arid Andes

Larval host plants can be the main resource underlying the distribution (including altitudinal ranges) of specialized phytophagous insects such as butterflies. Strymon flavaria (Ureta) (Lepidoptera: Lycaenidae: Theclinae: Eumaeini) is a little-known hairstreak endemic to the arid belt at about 3000–3500 m elevation on the western slopes of the Andes of northern Chile. The hemiparasitic shrub Krameria lappacea (Dombey) Burdet and Simpson is here recorded as the first host plant known for S. flavaria, representing also the first record of the association of a species of Strymon Hübner with the plant family Krameriaceae. Females of S. flavaria lay eggs on flower buds and unripe fruits of K. lappacea; the larva eats mostly these plant organs. Field observations revealed that S. flavaria is a host-specialist hairstreak, since females lay eggs only on K. lappacea. These findings suggest that the altitudinal range of S. flavaria could be the result of its specialized association with K. lappacea, as this shrub is also mostly restricted to elevations above 3000 m in the Andes of northern Chile. The implications of this finding for the ecology and conservation of S. flavaria are discussed.     

Effect of the loading rate on compressive properties of goose eggs

The resistance of goose (Anser anser f. domestica) eggs to damage was determined by measuring the average rupture force, specific deformation and rupture energy during their compression at different compression speeds (0.0167, 0.167, 0.334, 1.67, 6.68 and 13.36 mm/s). Eggs have been loaded between their poles (along X axis) and in the equator plane (Z axis). The greatest amount of force required to break the eggs was required when eggs were loaded along the X axis and the least compression force was required along the Z axis. This effect of the loading orientation can be described in terms of the eggshell contour curvature. The rate sensitivity of the eggshell rupture force is higher than that observed for the Japanese quail’s eggs.