A multiple-scale analysis of host plant selection in Lepidoptera

Lepidoptera play an important role in terrestrial ecosystems as pollinators, as components of the food chain and as indicators for healthy ecosystems due to their sensitivity to change. Heterogeneous landscapes with variability of topographical features, vegetation structure combined with food sources for all life stages are the basis for successful lepidopteran oviposition. A multiple-scale analysis is proposed to understand the hierarchical relationships between selected site to plant characteristics and oviposition preferences for the dingy skipper (Erynnis tages). To achieve this goal, factors driving oviposition at the plant and patch scale were identified and scale dependencies at the site scale were assessed. At the plant scale, tallest host plants were used for oviposition; however relative egg height upon each plant was similar in both host plant species [bird’s-foot trefoil (Lotus corniculatus) and horseshoe vetch (Hippocrepis comosa)]. The main factors preferred by E. tages in L. corniculatus patches were sward height and percent of bare ground, and in H. comosa host plant density patches. Selected patches had slopes of greater gradients (mostly facing south) than patches with no selected host plants. At the site scale, oviposition patches were clustered at small scales and oviposition sites were dispersed at larger scales. Our study suggests that oviposition selection in E. tages is a hierarchical process varying from the site to the plant scale. Our study provides empirical evidence useful to inform landscape management strategies. These can be expanded to assess larger scale vegetation and habitat suitability beyond individual sites for systematic conservation planning.     

Oviposition site selection of a threatened moth <Emphasis Type="Italic">Eriogaster catax</Emphasis> (Lepidoptera: Lasiocampidae) in agricultural landscape—implications for its conservation

Shrubby field margins and hedgerows play an important role for many species of flora and fauna in agro-ecosystems, including those which are rare or endangered. A characteristic species of these semi-natural habitats in agricultural landscape is the Eastern eggar, Eriogaster catax (Linnaeus, 1758): a moth species threatened in many European countries. In this study we investigated the oviposition preferences of E. catax at host plant level in agricultural landscape in south-western Poland, near of the northern limit of the species distribution in Europe. In total we surveyed 796 host plants. Altogether we found 439 eggs batches or caterpillar webs of E. catax on 373 occupied plants. Most of them (92.3%) were on blackthorn Prunus spinosa, which was preferred host plant next to pear Pyrus spp. However, the most important predictor of the presence of egg batches were microclimatic conditions. We recorded that females preferred host plants exposed to direct sunlight and with south and west-facing slopes for oviposition and avoided those inside of the dense shrub canopy. Eggs were laid more often on the trunk and on the upper half of the host plant, however there was a significant difference in vertical and horizontal position of egg batches in relation to the position of the plant. Inside of the shrub canopy, eggs were laid in higher parts of host plant and closer to the trunk, than at the edge and on solitary plants. Based on our observations the management strategies for the habitats of the species are proposed and discussed.     

Interpatch movement of the red milkweed beetle,Tetraopes tetraophthalmus: individual responses to patch size and isolation

Individual movement patterns and the effects of host plant patch size and isolation on patch occupancy were examined for red milkweed beetles, Tetraopes tetraophthalmus, residing in a heterogeneous landscape. Male beetles were found to move both more often and farther between host plant patches than female beetles, and this difference affected the patterns of patch occupancy observed. Overall, unoccupied milkweed patches were smaller and more isolated than patches occupied by beetles. Patches uninhabited by females tended to be more isolated, but not necessarily smaller, than patches with female beetles, indicating that females may be affected more by patch isolation than patch size. Presence of male beetles on patches showed a stronger response to patch size than to patch isolation. Differences in movement between males and females illustrate the need for demographically based dispersal data. Comparisons of Tetraopes interpatch movement patterns between landscapes composed of patches of different size revealed that landscapes with overall smaller patches may have greater rates of interpatch movement.     

Habitat quality matters for the distribution of an endangered leaf beetle and its egg parasitoid in a fragmented landscape

Fragmentation, deterioration, and loss of habitat patches threaten the survival of many insect species. Depending on their trophic level, species may be differently affected by these factors. However, studies investigating more than one trophic level on a landscape scale are still rare. In the present study we analyzed the effects of habitat size, isolation, and quality for the occurrence and population density of the endangered leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae) and its egg parasitoid, the hymenopteran wasp Foersterella reptans Nees (Hymenoptera: Tetracampidae). C. canaliculata is strictly monophagous on meadow sage (Salvia pratensis), while F. reptans can also parasitize other hosts. Both size and isolation of habitat patches strongly determined the occurrence of the beetle. However, population density increased to a much greater extent with increasing host plant density ( = habitat quality) than with habitat size. The occurrence probability of the egg parasitoid increased with increasing population density of C. canaliculata. In conclusion, although maintaining large, well-connected patches with high host plant density is surely the major conservation goal for the specialized herbivore C. canaliculata, also small patches with high host plant densities can support viable populations and should thus be conserved. The less specialized parasitoid F. reptans is more likely to be found on patches with high beetle density, while patch size and isolation seem to be less important.     

Host–parasitoid dynamics in a fragmented landscape: Holly trees,holly leaf miners and their parasitoids

Many species inhabit fragmented landscapes, where units of resource have a patchy spatial distribution. While numerous studies have investigated how the incidence and dynamics of individual species are affected by the spatial configuration and landscape context of habitat patches, fewer studies have investigated the dynamics of multiple interacting resource and consumer species in patchy landscapes. We describe a model system for investigating host–parasitoid dynamics in a patchy landscape: a network of 166 holly trees, a specialised herbivore of holly (the leaf miner, Phytomyza ilicis (Curtis, 1948)), and its suite of parasitoids. We documented patch occupancy by P. ilicis, its density within patches, and levels of parasitism over a 6-year period, and manipulated patch occupancy by creating artificially vacant habitat patches. Essentially all patches were occupied by the herbivore in each year, suggesting that metapopulation dynamics are unlikely to occur in this system. The main determinants of densities for P. ilicis and its parasitoids were resource availability (patch size and host density, respectively). While P. ilicis is apparently not restricted by the spatial distribution of resources, densities of its parasitoids showed a weaker positive relationship with host density in more isolated patches. In patches where local extinctions were generated experimentally, P. ilicis densities and levels of parasitism recovered to pre-manipulation levels within a single generation. Furthermore, patch isolation did not significantly affect re-colonisation by hosts or parasitoids. Analysing the data at a variety of spatial scales indicates that the balance between local demography and dispersal may vary depending on the scale at which patches are defined. Taken together, our results suggest that the host and its parasitoids have dispersal abilities that exceed typical inter-patch distances. Patch dynamics are thus largely governed by dispersal rather than within-patch demography, although the role of demography is higher in larger patches.     

Patch occupancy of two hemipterans sharing a common host plant

Aim Two hemipteran species were chosen as a study system for the comparative analysis of patch occupancy and spatial population structure of insects sharing a common host plant. This study tested whether (1) the incidence in the host plant patches differed between the two species, and (2) the two species exhibited a different spatial population structure, i.e. were they affected differentially by isolation and area of the host plant patches. Location The porphyry landscape north of Halle (Saale) in Germany comprising 506 patches of the host plant Brachypodium pinnatum. Methods The host plant patches were surveyed for the two hemipterans. To assess the influence of patch quality on species occurrence the patches were characterized by mean cover abundance of B. pinnatum, type of subsoil, slope, exposure, and shading. The spatial configuration of the patches was considered by patch area and isolation. The influence of the habitat factors and the spatial configuration on the occupancy of the two species was analysed by logistic regression. Results Adarrus multinotatus was found in 441 patches, while Neophilaenus albipennis was found in only 90 patches. While A. multinotatus showed virtually no relationship to the habitat factors, the occupancy of N. albipennis was influenced by subsoil type, cover abundance, and shading. The effects of area and isolation on occupancy of the patches also differed between the two species. The occupancy of N. albipennis was determined largely by area and isolation, whereas in A. multinotatus no considerable effect of spatial configuration was found. Main conclusions The study revealed a marked difference between the two hemipteran species in respect of spatial population structure. Adarrus multinotatus built up a ‘patchy population’, whereas N. albipennis showed a ‘metapopulation’ structure within the same set of patches in the same landscape. Spatial population structure was found to be not only a function of spatial configuration of habitat patches, but population structure differed between the habitat generalist A. multinotatus and the habitat specialist N. albipennis.     

Microhabitat selection in a grassland butterfly: a trade-off between microclimate and food availability

Understanding the factors that determine habitat quality is vital to ensuring appropriate habitat management. The main objective of this study was to assess the microhabitat preferences of egg-depositing females of the Grizzled Skipper (Pyrgus malvae) in calcareous grasslands of the Diemel Valley (Central Germany) for defining habitat quality. Based on this knowledge, we make management recommendations for the conservation of this threatened species. P. malvae generally preferred open and warm oviposition sites. However, there were considerable differences in the environmental conditions, depending on the selected host plant. On the small Potentilla tabernaemontani plants that grew in sparse vegetation with low-growing turf, mostly only one egg was found per plant. In contrast, occupied Agrimonia eupatoria host plants were larger and more prominent, regularly having more than one egg, and grew at sites with a taller and denser vegetation. The observed oviposition pattern reflects a trade-off between microclimate and food availability: Usually, occupied P. tabernaemontani plants grow under favourable microclimatic conditions. However, during hot years the risk of desiccation is high, leading to food shortage. In contrast, A. eupatoria generally provides more biomass, thrives on deeper soils and the vegetation has a cooler microclimate: hence, food shortage is somewhat unlikely. To meet the described habitat requirements of P. malvae, traditional rough grazing by sheep and goats seemed to be the most appropriate land management strategy. The re-introduction of coppicing in woodlands, particularly adjacent to calcareous grasslands, would also be beneficial.     

Effect of vegetation density, height, and connectivity on the oviposition pattern of the leaf beetle Galeruca tanaceti

Vegetation structure can profoundly influence patterns of abundance, distribution, and reproduction of herbivorous insects and their susceptibility to natural enemies. The three main structural traits of herbaceous vegetation are density, height, and connectivity. This study determined the herbivore response to each of these three parameters by analysing oviposition patterns in the field and studying the underlying mechanisms in laboratory bioassays. The generalist leaf beetle, Galeruca tanaceti L. (Coleoptera: Chrysomelidae), preferentially deposits its egg clutches on non‐host plants such as grasses. Earlier studies revealed that oviposition within structurally complex vegetation reduces the risk of egg parasitism. Consequently, leaf beetle females should prefer patches with dense, tall, or connected vegetation for oviposition in order to increase their reproductive success. In the present study, we tested the following three hypotheses on the effect of stem density, height, and connectivity on oviposition: (1) Within habitats, the number of egg clutches in areas with high stem densities is disproportionately higher than in low‐density areas. The number of egg clutches on (2) tall stems or (3) in vegetation with high connectivity is higher than expected for a random distribution. In the field, stem density and height were positively correlated with egg clutch presence. Moreover, a disproportionately high presence of egg clutches was determined in patches with high stem densities. Stem height had a positive influence on oviposition, also in a laboratory two‐choice bioassay, whereas stem density and connectivity did not affect oviposition preferences in the laboratory. Therefore, stem height and, potentially, density, but not connectivity, seem to trigger oviposition site selection of the herbivore. This study made evident that certain, but not all traits of the vegetation structure can impose a strong influence on oviposition patterns of herbivorous insects. The results were finally compared with data on the movement patterns of the specialised egg parasitoid of the herbivore in comparable types of vegetation structure.     

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.     

Oviposition preferences of a threatened butterfly Leptidea morsei (Lepidoptera: Pieridae) at the western border of its range

The oviposition preferences of Leptidea morsei at the western border of its range (SE Slovenia) were studied with the aim of integrating this information into species habitat management recommendations. The characteristics of egg-laying habitat were examined at the landscape, patch, plant and leaf levels. Because sightings of adults in the field were infrequent, intensive searching for previously laid eggs on Lathyrus niger plants was chosen as the survey method. The main oviposition habitats of L. morsei in Slovenia were found to be the forest edges and light stands of thermophilous oak, oak-hornbeam and mesic beech forests (landscape level). The egg-laying sites were L. niger plants growing individually or in small stands, and oriented mostly towards W, SW, and S. The oviposition plants were between 33 and 120 cm high, with those growing in half to full shade preferred. Eggs were deposited singly, mainly on the underside of leaflets of terminal leaves, located mostly on the top quarter of plants irrespective of their height and growth form. The most important factors affecting egg-placement are sun exposure and the size of L. niger stands at ovipositing sites (patch level), leaf exposure on the plant in vertical and horizontal directions (plant level), leaf age and the leaf parts where the risk of leaf and egg desiccation is lower (leaf level). Management strategies for the main egg-laying habitats of the species are proposed.     

Heterogeneity in patch quality buffers metapopulations from pathogen impacts

Many wildlife species persist on a network of ephemerally occupied habitat patches connected by dispersal. Provisioning of food and other resources for conservation management or recreation is frequently used to improve local habitat quality and attract wildlife. Resource improvement can also facilitate local pathogen transmission, but the landscape-level consequences of provisioning for pathogen spread and habitat occupancy are poorly understood. Here, we develop a simple metapopulation model to investigate how heterogeneity in patch quality resulting from resource improvement influences long-term metapopulation occupancy in the presence of a virulent pathogen. We derive expressions for equilibrium host–pathogen outcomes in terms of provisioning effects on individual patches (through decreased patch extinction rates) and at the landscape level (the fraction of high-quality, provisioned patches), and highlight two cases of practical concern. First, if occupancy in the unprovisioned metapopulation is sufficiently low, a local maximum in occupancy occurs for mixtures of high- and low-quality patches, such that further increasing the number of high-quality patches both lowers occupancy and allows pathogen invasion. Second, if the pathogen persists in the unprovisioned metapopulation, further provisioning can result in all patches becoming infected and in a global minimum in occupancy. This work highlights the need for more empirical research on landscape-level impacts of local resource provisioning on pathogen dynamics.     

Host plant use by the Heath fritillary butterfly, Melitaea athalia: plant habitat, species and chemistry

We present a study of habitat use, oviposition plant choice, and food plant suitability for the checkerspot butterfly Melitaea athalia Rottemburg (Lepidoptera: Nymphalidae) in Åland, Finland. We found that in Åland, unlike in the mainland of Finland and many parts of its range, M. athalia flies mainly in open meadows. When offered an array of plants in a large (32 × 26 m) field cage, they predominately oviposited upon Veronica chamaedrys L., V. spicata L. and Plantago lanceolata L. (Plantaginaceae), which grow in open meadows. The relative abundance of the butterfly in Åland, and its habitat and host plant use there, may reflect local adaptation to land use practices and geology that maintain clusters of small open meadows with little successional change. At the scale of a plant patch, preferred species were used as frequently in mixed species patches as in mono-specific patches, and more oviposition occurred in open than in grassy patches. All of the host plants used by M. athalia are defended by iridoid glycosides (IGs). However, oviposition choice among species and among individual plants within species was largely independent of IG concentration. This contrast with the more discerning congener, M. cinxia, supports the idea that host discrimination decreases with increasing host range. Finally, although the adult butterflies chose specific plant species for oviposition, as larvae they performed well on twelve out of thirteen species of plants, including both known hosts and related novel plants that occur in Åland, indicating a much wider range of larval food plant species than adult oviposition species.     

No Experimental Evidence for Host Ant Related Oviposition in a Parasitic Butterfly

The ability of adult butterflies of the genus Maculinea to locate their host ants prior to oviposition has been the subject of much discussion. We studied the egg laying behavior of the dusky large blue Maculinea nausithous whose larvae parasitize colonies of the ant Myrmica rubra. Flowerheads of the initial food plant were sprinkled with soil from ant nests, which contain chemicals involved in the nest recognition behavior of ants. The experiment was conducted to determine whether ant-released chemicals may act as oviposition cues and whether intraspecific competition for suitable plants may force female butterflies to alternative decisions. Host plant choice was not influenced by the presence of nest-derived host-ant cues. Density dependent shifts to less suitable host plants could not be ascertained nor changes in egg laying behavior across the flight period. The observed egg distribution could be primarily explained by host plant characteristics and environmental variability among sites. The result confirms the theory that host ant dependent oviposition appears to be a disadvantageous strategy in the face of resource limitation within ant colonies and the immobility of caterpillars.     

Behavioural plasticity in relation to egg and time limitation: the case of two fly species in the genus Anastrepha (Diptera: Tephritidae)

Reproductive opportunities in insects that deposit their eggs in discrete resource patches are frequently limited because the availability of oviposition substrates is often spatially and temporally restricted. Such environmental variability leads individuals to confront time‐ or egg‐limitation constraints. Additionally, species with different oviposition strategies (i.e. single egg layers vs clutch layers) commonly deal with different structural and ecological characteristics of larval host plants. To test the hypothesis that oviposition strategies such as laying eggs singly or in batches (clutches) are related to these constraints (i.e. egg vs time limitation), we compared the lifetime oviposition patterns of two closely related sympatric species of Anastrepha (Diptera: Tephritidae) with different oviposition strategies. We exposed five cohorts of A. obliqua and A. ludens females, over the course of their adult lifetimes, to three conditions of “habitat quality” (measured as host density per cage): unpredictable habitat quality (host density varied randomly from day to day between 1, 5, 15, 30 and 60 hosts/cage), low habitat quality (fixed density of one host/cage) and high habitat quality (fixed density of 60 hosts/cage).
Responses to host density conditions were strikingly different in the two species. (1) Frequency of host visits and oviposition events increased in A. obliqua but not in A. ludens when host densities increased. (2) Anastrepha ludens females accepted low quality hosts (i.e. fruits on which eggs had already been laid and were therefore partially covered with host marking pheromone) significantly more often than A. obliqua females did. (3) Females of A. obliqua adjusted their oviposition activity to variations in host density, whereas A. ludens females exhibited a constant oviposition pattern (i.e. did not respond to variations in host density). Based on the above, it is likely that in A. obliqua oviposition is governed by egg‐limitation and in A. ludens by time‐limitation constraints. We discuss the relationship between the oviposition strategies of each fly species and the fruiting phenology and density of their native host plants. We also address the possible influence of oogenesis modality and parasitism by braconid wasps in shaping oviposition behaviour in these insects.     

Combining larval habitat quality and metapopulation structure – the key for successful management of pre-alpine Euphydryas aurinia colonies

This study aims to analyse larval habitat preferences and landscape level population structure of the threatened Marsh Fritillary butterfly, Euphydryas aurinia, and discusses implications for the conservation and management of this strongly declining species in central Europe. Whereas current management strategies are mainly based on studies of habitat requirements of adult individuals, we intend to emphasise larval habitat quality and population processes at the landscape level as additional key factors. Microhabitat preference analysis of egg-laying females showed that eggs were predominantly laid on prominent large-sized host plant individuals. Additionally, when Succisa pratensis was used as a host plant (as opposed to Gentiana asclepiadea), host individuals in open vegetation structure were preferred. Optimal oviposition conditions were present in recently abandoned calcareous fen meadows and at the edges of such meadows currently in use. A two-year patch-occupancy study in the northern pre-alpine region of south-west Germany indicated that E. aurinia lives in a metapopulation. In a logistic-regression model, patch size, isolation, and habitat quality explained 82% of the observed patch-occupancy pattern in 2001. Our data suggest that a suitable conservation strategy must incorporate both the conservation of a network of suitable habitat patches, and efforts to maximise local habitat quality by ensuring that host plants can grow to a large size and are surrounded by sparse and low vegetation cover.     

The Duke of Burgundy butterfly and its dukedom: larval niche variation in <Emphasis Type="Italic">Hamearis lucina</Emphasis> across Central Europe

In order to improve our understanding of habitat preferences and optimal management of open woodland insects, we analyse patch occupancy and oviposition electivity of the endangered Duke of Burgundy butterfly, Hamearis lucina, in three regions across German habitat types. Some newly available forest clearings created by a severe winter storm in the Schönbuch region were colonised within 2 years, whereas some suitable patches remained unoccupied for several years. We discuss how small population sizes, limited patch connectivity, and habitat quality may contribute to such an intermediate colonisation power. Across study regions, we document differences in oviposition site electivity. On calcareous grassland in the Diemeltal, shaded Primula plants on western slopes were preferentially used, probably to avoid desiccation of the natal food plant. To the contrary, sun-exposed Primula stands were preferred in forest clearings in the Schönbuch and calcareous fens in the Allgäu. In these regions, the risk of desiccation is low, but the overall cool and moist microclimate conditions may hamper larval development in shaded conditions, favouring oviposition at sites with maximum exposition to solar radiation. Optimal management strategies depend on the abiotic and biotic conditions and conservation priorities for the given region and habitat type, and we suggest management regimes that promise to sustain suitable H. lucina habitat at a landscape-level.     

野外条件下广聚萤叶甲对豚草的产卵选择

为探究影响广聚萤叶甲Ophraella communa LeSage选择豚草产卵的环境因素, 在南京城郊于2009年6月23日至9月1日豚草生长季节, 跟踪调查了60个豚草斑块上落卵数量动态, 运用跨栏模型分析了影响叶甲产卵以及产卵数量的因素。结果表明： （1）产卵发生随季节延长、 豚草被取食程度增大、 株高增长、 暴露环境中斑块增大而增加, 但随遮蔽环境中豚草斑块增大、 开花、 开花后季节延长、 季节延长情况下被取食程度增大、 开花后被取食程度增大等因素而降低; （2）落卵数量随季节延长、 豚草被取食程度增大、 暴露环境、 开花期被取食程度增大、 开花期株高增长等因素而增加, 但随豚草开花、 季节延长情况下被取食程度增大而减小; （3）落卵量随季节延长而增大的幅度在豚草营养生长期明显大于开花期, 在暴露环境中随季节延长而呈明显直线增大, 但在遮阴环境中先直线增加, 于10周后趋缓。研究结果说明, 影响产卵发生与落卵数量的因素不完全一致, 产卵发生与落卵数量主要受因子之间互作的影响, 而较少受到单因素的独立影响。     

Temporal patch occupancy dynamics of the Siberian flying squirrel in a boreal forest landscape

Ability to predict species distribution in a landscape is of crucial importance for natural resource management and species conservation. Therefore, the understanding of species habitat requirements and spatio-temporal dynamics in occurrence is needed. We examined patch occupancy patterns of the Siberian flying squirrel Pteromys volans in northern Finland across a seven year study period. Forest patches dominated by mature spruce ( Picea abies ) in a study area (375 km²) were surveyed to monitor the presence or absence of the flying squirrel. The patch occupancy pattern was dynamic: about half of the habitat patches were occupied at least once during the study period and more patches were colonised than were abandoned. Patches that were continuously occupied (i.e. occupied during all sample periods) were typically of high quality (based on habitat and landscape characteristics), continuously unoccupied patches were usually of low quality, and intermediate quality patches were occupied intermittently. The variables explaining patch occupancy were similar each year, and a statistical model based on data from the year 2000 also predicted occupancy in 2004 with similar accuracy. However, data from a single survey were inadequate for identifying patches used intermittently by flying squirrels. Despite inconsistent occupancy, these patches may be important for the local persistence of flying squirrels. The dynamic occupancy pattern may thus affect estimates of suitable habitat area and identification of functional patch networks for landscape planning. These results emphasise the need for follow-up studies to better understand population patterns and processes in time.     

Management of calcareous grasslands for Nickerl’s fritillary (<Emphasis Type="Italic">Melitaea aurelia</Emphasis>) has to consider habitat requirements of the immature stages,isolation, and patch area

We analysed the habitat preferences of adult stages and oviposition electivity of Melitaea aurelia in calcareous grasslands in the Diemel Valley (central Germany) to assess the key factors for successful management. Egg-laying and adult habitats of M. aurelia were more or less congruent. Oviposition electivity at the host plant (Plantago media) was best explained by a combination of host plant quantity and vegetation structure. Habitat quality, isolation and patch area explained 86% of the current patch occupancy of M. aurelia. With M. aurelia preferentially inhabiting transitional vegetation types, management requires a balance between abandonment and disturbance. Disturbances provide open soil that facilitates germination of the host plant Plantago media. On the other hand, immature and adult stages of M. aurelia perform best on calcareous grasslands with a high amount of host plants and low disturbance intensity. Traditional rough grazing regimes seem to be the most favourable tool for developing the necessary spatial and temporal heterogeneity in patches. The best results may be achieved by rotational grazing where only a subset of inhabited patches is grazed intensively each year. Our analysis of patch occupancy indicates that it would be desirable to restore patches in close proximity to occupied sites.     

Multitrophic influences on egg distribution in a specialized leaf beetle at multiple spatial scales

Egg distribution in herbivorous beetles can be affected by bottom-up (host plant), and by top-down factors (parasitoids and predators), as well as by other habitat parameters. The importance of bottom-up and top-down effects may change with spatial scale.

In this study, we investigated the influence of host plant factors and habitat structure on egg distribution in the leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae), a monophagous herbivore on Salvia pratensis L. (Lamiales: Lamiaceae), on four spatial scales: individual host plant, microhabitat, macrohabitat, and landscape. At the individual host plant scale we studied the correlation between egg clutch incidence and plant size and quality. On all other scales we analyzed the relationship between the egg clutch incidence of C. canaliculata and host plant percentage cover, host plant density, and the surrounding vegetation structure. Vegetation structure was examined as herbivores might escape egg parasitism by depositing their eggs on sites with vegetation factors unfavorable for host searching parasitoids.

The probability that egg clutches of C. canaliculata were present increased with an increasing size, percentage cover, and density of the host plant on three of the four spatial scales: individual host plant, microhabitat, and macrohabitat. There was no correlation between vegetation structure and egg clutch occurrence or parasitism on any spatial scale. A high percentage of egg clutches (38–56%) was parasitized by Foersterella reptans Nees (Hymenoptera: Tetracampidae), the only egg parasitoid, but there was no relationship between egg parasitism and the spatial distribution of egg clutches of C. canaliculata on any of the spatial scales investigated. However, we also discuss results from a further study, which revealed top-down effects on the larval stage.