Responses of plant, insect and spider biodiversity to local and landscape scale management intensity in cereal crops and grasslands

The aim of this study was to determine the relative effects of landscape scale management intensity, agroecosystem type, local management intensity and edges on diversity patterns of functional groups of plants, carabid beetles, spiders and grasshoppers. Nine landscapes were selected differing in percent intensively used agricultural area (IAA), each with a pair of organic and conventional winter wheat fields and a pair of organic and conventional mown meadows. Within fields, plants were surveyed in the edge and in the interior. Carabid beetles and spiders were captured by funnel traps, while grasshoppers were sweep-netted in the meadows. Diversity patterns of study organisms were affected both by local variables (local management, agroecosystem type and within-field position) and by landscape scale management intensity. Species richness of grasses, presumably because of sowing low-diversity mixtures, and hunting spiders decreased with percent cover of IAA. Meadows differed from wheat fields in that they had higher species richness of forbs and grasses, as well as higher densities of hunting spiders. In contrast, more carabid individuals, especially of non-carnivore species, were captured in wheat fields. In field edges with their reduced management intensity and increased immigration, species richness of plants, carabids and spiders was higher than in the interiors regardless of agroecosystem type and management. Organic management enhanced forb richness and cover in both agroecosystem types. Organic management also increased grass cover in wheat fields, but not in meadows, and promoted species richness of non-carnivore carabids and hunting spiders, but not grasshoppers. The results show that agri-environmental management needs to be targeted to the agroecosystem's field size, because higher edge area led to higher species richness. Organic management affected several functional groups positively (forbs, non-carnivore carabids, hunting spiders), while lower landscape scale management intensity only increased species richness of grasses and spiders. The great differences in responses of functional groups to local cereal and grassland as well as landscape management suggest implementing more scale and group specific targets for agri-environmental schemes to improve their efficiency.     

Delayed colonisation of arable fields by spring breeding ground beetles (Coleoptera: Carabidae) in landscapes with a high availability of hibernation sites

The colonisation of winter barley fields by spring breeding carabids and its temporal modulation by the amount of potential hibernation sites was studied. Species richness of carabids was lower in landscapes with high length of boundaries and a high amount of non-cropped open habitats during early stages of the beetles’ colonisation of arable fields. Species number of beetles with high dispersal potential responded to this landscape features at coarse spatial scales whereas beetles with low dispersal potential responded to intermediate scales. However, the negative impact of potential hibernation sites on colonisation diminished in later sampling phases. The patterns observed may be explained by both overwintering in arable soils in less complex landscapes and delayed colonisation in more complex landscapes. The seasonal patterns of landscape control suggest a need to account for temporal dynamics in interactions between species or functional groups and landscape properties. A high temporal resolution is needed in studies that focus on ecosystem function and services in agricultural landscapes, as direction of effect (positive/negative) of management on animal communities may change across spatial scales and within short time periods.     

Impact of agricultural management on carabid communities and weed seed predation

This study evaluated the relationship between diversity and activity-density of carabid beetles and invertebrate weed seed predation in conventional, no-till, and organic management systems in the Midwest USA. Carabid beetles were sampled with pitfall traps and invertebrate seed predation rates of fall panicum and common lambsquarters were assayed with exclosure cages. Total carabid activity-density was over two times higher in the conventional systems compared to the no-till and organic management systems. In contrast, activity-densities of seed-predating carabid species were over three times higher in the no-till compared to the conventional and organic systems. Carabid diversity was higher in the no-till and organic systems compared to the conventional system, and a multivariate analysis showed that carabid community structure was distinct among the three systems. Predation of fall panicum and common lambsquarters seeds was often over two times higher in the no-till compared to the conventional and organic systems, and there was a strong correlation (r > 0.94) between seed removal rates and the total number of carabid seed predators captured in each system.     

Comparison of riparian plant communities under four land management systems in southwestern Wisconsin

Riparian plant community composition is influenced by moisture, erosion, original native plant communities, and current and past land use. This study compared riparian plant communities under four types of management: woody buffer strip, grassy buffer strip, rotational grazing, and continuous grazing. Study sites were located along spring-fed streams in the unglaciated region of southwestern Wisconsin, USA. At each site, plant community surveys were conducted using a point transect method. Among the treatments, woody buffer strips, rotationally grazed and continuously grazed riparian areas had greater plant species richness than grassy buffer strips, and woody buffer strips had the greatest native plant species richness. Reed canarygrass (Phalaris arundinacea L.) was prevalent in grassy buffer strips (44% of all observations), common in woody buffer strips (15%), and rare in sites that were rotationally or continuously grazed (3 and 5%, respectively). Pasture sites had greater proportions of native grasses and grass relatives and moderate levels of overall native species richness. Considered a water quality best management practice, well-managed rotational grazing may be a reasonable alternative to buffer strips which can contribute to protection and enhancement of native vegetation biodiversity.     

Density and nutritional condition of carabid beetles in wildflower areas of different age

Density and nutritional condition of the five carabid beetle species Agonum mülleri, Anchomenus dorsalis, Anisodactylus binotatus, Pterostichus vernalis and Poecilus cupreus was studied. Carabids were caught in sixteen 1–4-year-old wildflower areas, identified and counted, their nutritional condition was calculated and calculations were related to habitat parameters (i.e. wildflower area age, vegetation cover, soil water content, habitat size, surrounding landscape composition) and sex.Whereas the number of Anisodactylus binotatus caught responded positively to succession in wildflower areas, A. mülleri numbers tended to decrease and the other three species were unaffected. In a stepwise multiple regression species affiliation, sex and age of wildflower area explained together 98.5% of the variance in the pooled nutritional condition of Anchomenus dorsalis, Anisodactylus binotatus, A. mülleri and P. vernalis. Age of wildflower area alone explained 61.7% of the variance and age increased the nutritional condition of carabids. Correspondingly, nutritional condition of Poecilus cupreus was positively correlated with the age of wildflower area. Overall, the nutritional condition of the carabid assemblage studied increased from the first to the second year and then remained stable. To keep the proportion of 1-year-old wildflower areas low, wildflower areas should therefore be maintained for several years to improve the life conditions of carabid beetles and to enhance their beneficial effects in agroecosystems.     

Grassy field margins and arthropod diversity: a case study on ground beetles and spiders in eastern Austria (Coleoptera: Carabidae; Arachnida: Aranei, Opiliones)

In 1983, the species composition and diversity of carabids, spiders and harvestmen was investigated by pitfall traps in a grassy field margin and in an adjacent, biologically farmed wheat field at Obere Lobau, Vienna, Austria.The carabid fauna of both habitats was diverse; it showed a high degree of similarity, the abundant species in both habitats being represented by typical field species. Only Metophonus spp. predominantly lived in the margin. Numerous species of different habitat origin occurred in the margin at low abundances indicating transitional stay rather than permanent populations.In contrast, the spider fauna showed a low degree of similarity, with few agrobiotic species strongly dominating field fauna. In the field margin, a rich spider fauna containing rare elements indicates the general importance of grassy margins for nature conservation. The apparently low faunal exchange of spiders between the two habitat types is discussed. Harvestmen occurred mainly in the field margin; only two thermophilic species invaded the field.Seasonal fluctuations in the trapping numbers of Bembidion lampros, Platynus dorsalis and Brachinus explodens are interpreted as indicating migrations between field margin and field. Poecilus cupreus and certain other field species, however, apparently spend their entire life cycle in the field. The possible function of field margins for restoration of carabid losses due to impacts of intensive cultivation is discussed.It is concluded that both grassy field margins and sustainable farming methods, in particular biological farming, are necessary to maintain a diverse predatory arthropod fauna in agroecosystems.     

Prison construction and guarding behaviour by European honeybees is dependent on inmate small hive beetle density

Increasing small hive beetle (Aethina tumida Murray) density changes prison construction and guarding behaviour in European honeybees (Apis mellifera L.). These changes include more guard bees per imprisoned beetle and the construction of more beetle prisons at the higher beetle density. Despite this, the number of beetles per prison (inmate density) did not change. Beetles solicited food more actively at the higher density and at night. In response, guard bees increased their aggressive behaviour towards beetle prisoners but did not feed beetles more at the higher density. Only 5% of all beetles were found among the combs at the low density but this percentage increased five-fold at the higher one. Successful comb infiltration (and thus reproduction) by beetles is a possible explanation for the significant damage beetles cause to European honeybee colonies in the USA.     

Use of pheromone traps to suppress populations of Scolytus multistriatus (Marsham) (Coleoptera: Scolytidae) in three isolated communities of elms

The technique of mass trapping as a pest management tool was evaluated for its potential in reducing populations of Scolytus multistriatus. More than two million beetles were caught on traps baited with a synthetic attractant composed of both aggregation pheromones and host tree volatiles. Trends in trap catches over a period of 4 years (1976–1979) in three small isolated towns of similar size and with a similar number of elms, suggest that there was little or no reduction of the beetle population. Patterns in the number of beetles captured and the dispersion of trap catches indicate that pheromonebaited traps may be deployed to help identify local infestations and to focus sanitation efforts.     

Design and use of a time-sorting pitfall trap for predatory arthropods

An electronically-controlled time-sorting pitfall trap was designed which improves upon earlier models in having a 24-h rotation time, and in being capable of running continuously for at least one week without attention. Four prototypes were built, using readily available, inexpensive components, and used from June to October, 1995, to collect carabid beetles and linyphiid spiders in clover-undersown and weeded cabbages. The traps operated without malfunction during the hottest weather ever recorded in Aberdeen, when soil surface temperatures exceeded 40°C, and also during periods of torrential rain. Three spider and two carabid species dominated the catch, their nocturnal and diurnal patterns of activity being in general agreement with other published data. Some predators were apparently able to adapt to the contrasting vegetation in the two treatments, two species of Erigone spiders being nocturnal in weeded cabbages but tending to be diurnal in undersown crops.     

Social encapsulation of beetle parasites by Cape honeybee colonies (Apis mellifera capensis Esch.)

Worker honeybees (Apis mellifera capensis) encapsulate the small hive beetle (Aethina tumida), a nest parasite, in propolis (tree resin collected by the bees). The encapsulation process lasts 1-4 days and the bees have a sophisticated guarding strategy for limiting the escape of beetles during encapsulation. Some encapsulated beetles died (4.9%) and a few escaped (1.6%). Encapsulation has probably evolved because the small hive beetle cannot easily be killed by the bees due to its hard exoskeleton and defensive behaviour.     

Matching innovations with potential users, a case study of potato IPM practices

Through interviews and mail surveys, Ohio potato (Solanum tuberosum) growers were surveyed concerning their willingness to adopt three cultural controls being developed for management of the Colorado potato beetle, Leptinotarsa decemlineata (Say), which had become increasingly difficult to control. The principal objective of the research was to obtain a deeper understanding of how farmers make decisions to adopt or not adopt new techniques and how these decisions are related to farmers' educational backgrounds and the way they currently manage their farms. The three cultural management options emphasized ecological approaches to habitat management rather than ecologically disruptive chemical control: (1) using host plants in overwintering sites to slow the spring movement of the beetles to potato fields, (2) using larger potato plants at field borders than in the center of the field as a spring trap crop, and (3) concentrating beetles in relatively small undefoliated areas to be killed with heat in late summer. Most respondents would not adopt these alternative pest management strategies, mainly because they perceived an unsatisfactory trade-off between logistic difficulties and expenses and population suppression they would achieve compared with traditional chemical control measures. Education correlated positively and experience in farming negatively with the willingness to try new pest management techniques. Additionally, growers responding more negatively to questions regarding integrated pest management strategies were more willing to experiment with the alternative techniques, a result attributed to either the survey design or an indication of experience with the more intensive management effort required for integrated pest management. This profile of potential innovators should be used by researchers to establish partnerships with farmers that could assist with both the research and development of new farming techniques and the adoption of successful systems by other farmers.     

Orange/lemon-scented beetles: opposite enantiomers of limonene as major constituents in the defensive secretion of related carabids

The major constituent in the pygidial gland defensive fluid of the carabid beetle Ardistomis schaumii is (R)-(+)-limonene, whereas that of Semiardistomis puncticollis is (S)-(−)-limonene. This was an unanticipated result, since it is not very common to find the opposite enantiomers of the same compound among the secondary metabolites of related species. Moreover, the glandular liquid of A. schaumii contains 1,8-cineole, and that of S. puncticollis has β-pinene, β-phellandrene, sabinene, and p-cymene. Of about 500 carabid species that have been chemically investigated, this is the first report of the presence of such complex mixtures of monoterpenes in their defensive secretions.     

Survey of current crop management practices in a mixed-ricefield landscape,Mekong Delta,Vietnam — potential of habitat manipulation for improved control of citrus leafminer and citrus red mite

In the Mekong Delta, Vietnam, the citrus leafminer Phyllocnistis citrella (CLM) and the citrus red mite Panonychus citri are major pests in both sweet orange (Citrus sinensis) and Tieu mandarin (C. reticulata). Survey data indicate that these pest problems might be aggravated after farmers have completely destroyed the weed flora in their orchard. As citrus farmers only perceive the larger predators such as the weaver ant Oecophylla smaragdina and spiders, and have no idea about the existence of predatory mites or parasitoids, they do not know about potential positive attributes of weeds in pest management which sustain populations of natural enemies and their alternative food. IPM training programmes could use the weaver ant as an introduction to educating farmers about predatory mites and parasitoids, and should likewise emphasise the importance of beneficial asteraceous weeds such as Ageratum conyzoides. Non-crop trees such as Spondias dulcis, Mangifera indica, Eucalyptus tereticornis and Ceiba pentandra are commonly known to offer good refuge for the weaver ant. These trees should be further studied for their temporal contribution as food resource for other natural enemies of CLM and mites. Small adjustments of current weed management techniques are suggested to improve availability of pollen and nectar for beneficials at crucial moments in the cropping season, with due respect to implications at the landscape level.     

Molecular evidence of facultative intraguild predation by Monochamus titillator larvae (Coleoptera: Cerambycidae) on members of the southern pine beetle guild

The southern pine bark beetle guild (SPBG) is arguably the most destructive group of forest insects in the southeastern USA. This guild contains five species of bark beetles (Coleoptera: Curculionidae: Scolytinae): Dendroctonus frontalis, Dendroctonus terebrans, Ips avulsus, Ips calligraphus, and Ips grandicollis. A diverse community of illicit receivers is attracted to pheromones emitted by the SPBG, including the woodborers Monochamus carolinensis and Monochamus titillator (Coleoptera: Cerambycidae). These woodborers have been traditionally classified as resource competitors; however, laboratory assays suggest that larval M. carolinensis may be facultative intraguild predators of SPBG larvae. This study used polymerase chain reaction (PCR)-based molecular gut content analyses to characterize subcortical interactions between M. titillator and members of the SPBG. The half-lives of SPBG DNA were estimated in the laboratory prior to examining these interactions in the field. A total of 271 field-collected M. titillator larvae were analyzed and 26 (9.6?%) tested positive for DNA of members of the SPBG. Of these larvae, 25 (96.2?%) tested positive for I. grandicollis and one (3.8?%) for I. calligraphus. Failure to detect D. terebrans and D. frontalis was likely due to their absence in the field. I. avulsus was present, but primers developed using adult tissues failed to amplify larval tissue. Results from this study support the hypothesis that larval Monochamus spp. are facultative intraguild predators of bark beetle larvae. Additionally, this study demonstrates the capabilities of PCR in elucidating the interactions of cryptic forest insects and provides a tool to better understand mechanisms driving southern pine beetle guild population fluctuations.     

Oviposition by small hive beetles elicits hygienic responses from Cape honeybees

Two novel behaviours, both adaptations of small hive beetles (Aethina tumida Murray) and Cape honeybees (Apis mellifera capensis Esch.), are described. Beetles puncture the sides of empty cells and oviposit under the pupae in adjoining cells. However, bees detect this ruse and remove infested brood (hygienic behaviour), even under such well-disguised conditions. Indeed, bees removed 91% of treatment brood (brood cells with punctured walls caused by beetles) but only 2% of control brood (brood not exposed to beetles). Only 91% of treatment brood actually contained beetle eggs; the data therefore suggest that bees remove only that brood containing beetle eggs and leave uninfected brood alone, even if beetles have accessed (but not oviposited on) the brood. Although this unique oviposition strategy by beetles appears both elusive and adaptive, Cape honeybees are able to detect and remove virtually all of the infested brood.     

Beetle adhesive hairs differ in stiffness and stickiness: in vivo adhesion measurements on individual setae

Leaf beetles are able to climb on smooth and rough surfaces using arrays of micron-sized adhesive hairs (setae) of varying morphology. We report the first in vivo adhesive force measurements of individual setae in the beetle Gastrophysa viridula, using a smooth polystyrene substrate attached to a glass capillary micro-cantilever. The beetles possess three distinct adhesive pads on each leg which differ in function and setal morphology. Visualisation of pull-offs allowed forces to be measured for each tarsal hair type. Male discoidal hairs adhered with the highest forces (919?±?104?nN, mean?±?SE), followed by spatulate (582?±?59?nN) and pointed (127?±?19?nN) hairs. Discoidal hairs were stiffer in the normal direction (0.693?±?0.111?N?m^?1) than spatulate (0.364?±?0.039?N?m^?1) or pointed (0.192?±?0.044?N?m^?1) hairs. The greater adhesion on smooth surfaces and the higher stability of discoidal hairs help male beetles to achieve strong adhesion on the elytra of females during copulation. A comparison of pull-off forces measured for single setae and whole pads (arrays) revealed comparable levels of adhesive stress. This suggests that beetles are able to achieve equal load sharing across their adhesive pads so that detachment through peeling is prevented.     

Skimming behaviour and spreading potential of Stenus species and Dianous coerulescens (Coleoptera: Staphylinidae)

Rove beetles of the genus Stenus Latreille and the genus Dianous Leach possess pygidial glands containing a multifunctional secretion of piperidine and pyridine-derived alkaloids as well as several terpenes. One important character of this secretion is the spreading potential of its different compounds, stenusine, norstenusine, 3-(2-methyl-1-butenyl)pyridine, cicindeloine, α-pinene, 1,8-cineole and 6-methyl-5-heptene-2-one. The individual secretion composition enables the beetles to skim rapidly and far over the water surface, even when just a small amount of secretion is emitted. Ethological investigations of several Stenus species revealed that the skimming ability, skimming velocity and the skimming behaviour differ between the Stenus species. These differences can be linked to varied habitat claims and secretion saving mechanisms. By means of tensiometer measurements using the pendant drop method, the spreading pressure of all secretion constituents as well as some naturally identical beetle secretions on the water surface could be established. The compound 3-(2-methyl-1-butenyl)pyridine excelled stenusine believed to date to be mainly responsible for skimming relating to its surface activity. The naturally identical secretions are not subject to synergistic effects of the single compounds concerning the spreading potential. Furthermore, evolutionary aspects of the Steninae’s pygidial gland secretion are discussed.     

Diet-dependent heat emission reveals costs of post-diapause recovery from different nutritional sources in a carnivorous beetle

Restoration of fat stores is metabolic first priority for many insects that emerge from hibernation with depleted fat bodies. To some extent, the animals must be flexible and use whatever foods available irrespective of their nutrient composition. Previously, the carabid beetles Anchomenus dorsalis have been found to refill their fat stores to the same extent over 9 days irrespective of the nutrient composition of their food. However, a higher cost of fat deposition when the food was rich in sugar or protein rather than lipid was indicated by higher total energy consumption. Here, we test the hypothesis of increased metabolic costs of building fat stores from sugar- or protein-rich food than from lipid-rich food by microcalorimetry. We measured the heat emitted from beetles that had fed on sugar-, protein-, or lipid-rich food for 0 (common control), 2, 5, or 10 days. As predicted, heat emission was increased in beetles getting sugar- and protein-rich food compared with those getting lipid-rich food. However, we did not confirm the beetles’ ability to rebuild fat stores from protein-rich food; instead, they increased in lean mass. Overall, sugar-rich food seems to be optimal for post-winter recovery, because it is better than lipid-rich food that allows concurrent rebuilding of fat stores and lean mass, which may benefit preparation for spring migration and reproduction. We propose that overwintered fruits may be highly preferred post-diapause food for these otherwise mostly carnivorous beetles.     

Avoidance of nonhost plants by a bark beetle, Pityogenes bidentatus, in a forest of odors

The bark beetle, Pityogenes bidentatus (Coleoptera: Scolytidae), searches in mixed conifer and deciduous forests of northern Europe for suitable branches of its host, Scots pine (Pinus sylvestris). We tested whether odors from several diverse nonhost trees and plants common in the habitat (e.g., mountain ash, Sorbus aucuparia; oak, Quercus robur; alder buckthorn, Frangula alnus; blueberry, Vaccinium myrtillus; raspberry, Rubus idaeus; and grass, Deschampsia flexuosa) would reduce the attraction of the bark beetle to traps releasing its aggregation pheromone components in the field. Volatiles from the leaves or bark of each of these plants significantly reduced the attraction of the beetles to their pheromone. Odors collected from these nonhosts and analyzed by GC/MS contained monoterpenes, sesquiterpenes, and green-leaf alcohols, several of which (e.g., 1-octene-3-ol and -caryophyllene) reduced the attraction to pheromone in the field and elicited electroantennographic responses. In the laboratory, reproduction by the beetle was marginal in nonhost Norway spruce, Picea abies, and was absent in the other nonhost trees. Olfactory avoidance of unsuitable nonhosts may have evolved due to advantages in avoiding mistakes during host selection.