Tree species richness and density affect parasitoid diversity in cacao agroforestry

In Brazil, cacao is mostly planted beneath shade trees. The diversity of shade trees varies from monospecific to highly diverse canopies, characteristic of pristine Atlantic Forest. This study evaluates the relationships between family richness of Hymenoptera-Parasitica and Chrysidoidea, and tree species richness and density, the species richness of herbaceous understorey, and the area and age of the cacao agroforestry system. We sampled 16 cacao agroforestry systems, with canopy diversity ranging from one to 22 tree species per hectare, in three seasons: summer (March), winter (August) and spring (November). Parasitoids were sampled using eight Malaise-Townes traps per site. Tree species richness and density were enumerated within 1 ha at each site, and herbaceous plant species richness was calculated in eight 1 m² plots, within the hectare. The number of parasitoid families increased with tree species richness and density in spring and summer, but decreased in winter. Neither species richness of herbaceous plants nor area and age of the system affected parasitoid family richness. We suggest that the increase of parasitoid diversity with tree species richness and density in warmer seasons reflects increasing heterogeneity and availability of resources. The decrease in parasitoid family number with tree density in winter may be due to local impoverishment of resources, leading to parasitoid emigration to neighbouring forest remnants. This result implies that a higher diversity of shade trees will help to maintain high parasitoid levels and, in consequence, higher levels of natural enemies of cacao pests, particularly in the warmer seasons. This prediction is borne out in the experience of cacao producers. The proper management of shade tree diversity will play a vital role in maintaining the sustainability of cacao agroforestry production systems in the tropics and, concurrently, will maintain high biodiversity values in these locations.     

Spectral cathodoluminescence analysis of hymenopteran mandibles with different levels of zinc enrichment in their teeth

The inclusion of Zn in insect mandibles affects their hardness and is functional to their use during feeding or reproducing. However, little is known on the chemical/structural base of Zn enrichment. Here, we found that cathodoluminescence (CL) technique revealed two different types of CL spectra in the mandibles of Hymenoptera, depending on the Zn enrichment level assessed by Energy Dispersive X-ray Spectroscopy (EDS). Individuals having negligible traces to low % of Zn in their mandible teeth (≤3 wt%) presented a wide band of luminescence in the visible range which resembled those observed in the CC structures of graphite. This spectrum is probably characteristic for un-enriched cuticle, since it did not differ from those obtained from the Zn-lacking inner part of mandibles. Individuals with moderate to high % of Zn in their mandible teeth (≥7 wt%), instead, presented additional CL peaks in the ultraviolet range. Comparisons with different minerals of Zn suggest that these peaks could be related with OZnO bonds, with hydroxyl groups and with zinc-chlorine links (in agreement with Cl high levels detected by the EDS). Being a non-destructive technique, CL allows large comparative studies of the chemistry of metal-enriched insect cuticle even using unique specimens, such as those deposited in Natural History Museums.     

Alternative mating tactics and extreme male dimorphism in fig wasps

The dimorphisms in morphology and behaviour of male fig wasps are among the most extreme in the animal kingdom, and offer excellent opportunities to test the predictions of certain sexual selection models. Winged males resemble their conspecific females closely, but wingless males are so divergent in form that they have repeatedly been classified into different taxa. Wingless males mate within their natal fig fruits, whereas winged males disperse to mate. Individual species may have winged males, wingless males or both morphs. A key hypothesis proposes that sexual selection on male mating opportunities favours winged males in species with small broods and wingless males in species with large broods. Using data from 114 species in 33 genera, we show that both simple and formal comparative analyses support the correlated evolution of large brood size and male winglessness. Theoretical models further predict that, in male dimorphic species, the proportion of winged males should equal (in cases without local mate competition) or exceed (in cases with local mate competition) the proportion of females developing in fig fruits without wingless males. These predictions are met by eight out of nine male dimorphic species studied. Taken together, the patterns across all species, and between different male dimorphic species, strongly support sexual selection on mating opportunities as the major determinant of male morph ratios in fig wasps.     

Competition for honeydew between two social wasps in South Island beech forests,New Zealand

Summary Honeydew is a sugary secretion of beech scale insects (Ultracoelostoma spp.). Two introduced species of wasps forage on the sugar droplets in New Zealand beech forests. We hypothesize that competition between them may explain whyVespula germanica has become locally extinct in beech (Nothofagus) forest invaded byV. vulgaris. Changes in behaviour of the two wasp species in response to changes in the honeydew resource were monitored. Foraging and rainfall separately and together reduced the honeydew standing crop. In response to the standing crop decreasing, more wasps were found on honeydew trees, they became less active, spent more time lapping the tree surface, and ingested drops of honeydew at a slower rate.V. vulgaris was more active, and obtained drops and energy at a higher rate thanV. germanica. These behavioural differences may lead to competitive advantages affecting queen size and possibly survival.     

Proteomic analysis of the venom of the social wasp Apoica pallens (Hymenoptera: Vespidae)

Wasps are a diverse group of insects that possess a sting apparatus associated with a venom gland, which is used for predation and colony defense. The biochemistry of Hymenoptera venom has been evaluated in relation to allergy and immunology, and proteomics has been shown to be a powerful tool for the identification of compounds with pharmacological potential. Data on wasps venom the of genus Apoica are scarce, so the objective of the present work was to identify the venom proteins of the eusocial wasp Apoica pallens, as a first step towards further investigation of applied uses of the venom and its protein constituents. The venom proteins were separated by two-dimensional gel electrophoresis, followed by MALDI-TOF/TOF mass spectrometry. A total of 259 spots were detected, with molecular weights from 4.9 to 141 kDa. Thirty of these proteins were identified and classified into eight functional categories: allergen, enzyme, metabolism, structural, environmental response, proteoglycan, active in DNA and RNA, and unknown function. Due to the few available proteomic data for wasp venom, many proteins could not be identified, which makes studies with proteomic analysis of Hymenoptera venom even more important.     

Conflict of interest in a mutualism: documenting the elusive fig wasp-seed trade-off

The generally accepted view that mutualisms represent reciprocal exploitations implies a greater or lesser degree of inherent tension between the partners. This view emphasizes the importance of identifying conflicts of interest between the partners, and then attempting to quantify the effects of factors that influence costs and benefits to each. The natural history of the speciose fig-fig wasp pollination mutualisms permits such measurements. However, previous attempts to document the presumed tensions, which are expected to result in a negative relationship between the production of viable seeds and pollinator wasp offspring, have met with mixed results, casting doubt on the existence of the conflict. Here, we present hierarchical analyses of 929 fruits sampled from 30 crops representing nine species of monoecious New World figs. These analyses control for the confounding influences of variation in (1) pollination intensity (numbers of foundress pollinators); (2) flower number per fruit; and (3) the proportion of those flowers that develop, on seed and wasp production, both among and within crops. We thereby show that a negative relationship between the production of viable seeds and wasps is, in fact, ubiquitous, thus documenting this underlying tension inherent in the mutualism. We suggest that complex interactions of variables that influence costs and benefits are likely to be a general property of most mutualistic systems. <br>     

Active vision in insects: an analysis of object-directed zig-zag flights in wasps (Odynerus spinipes , Eumenidae)

Ground-nesting wasps (Odynerus spinipes, Eumenidae) perform characteristic zig-zag flight manoeuvres when they encounter a novel object in the vicinity of their nests. We analysed flight parameters and flight control mechanisms and reconstructed the optical flow fields which the wasps generate by these flight manoeuvres. During zig-zag flights, the wasps move sideways and turn to keep the object in their frontal visual field. Their turning speed is controlled by the relative motion between object and background. We find that the wasps adjust their rotational and translational speed in such a way as to produce a specific vortex field of image motion that is centred on the novel object. As a result, differential image motion and changes in the direction of motion vectors are maximal in the vicinity and at the edges of the object. Zig-zag flights thus seem to be a `depth from motion' procedure for the extraction of object-related depth information. Accepted: 31 August 1997     

Insect pollinator diversity in four forested ecosystems of southern Punjab,Pakistan

This study investigated pollinator assemblage diversity and richness in four forested ecosystems of southern Punjab, Pakistan, with different landscape types. Pirowal is situated in the plains of irrigated Punjab, Lal Suhanra is part of a sandy desert ecosystem, Ghazi Ghat is part of the Indus River delta, and Fort Munro is located in dry hilly mountains. A yearlong survey of pollinator populations was carried out in these four forested ecosystems from January to December of 2010. Fortnightly hand netting was performed for collecting flower-visiting insects whereas, pan traps of three colors (white, blue, and yellow) were deployed for collecting the data. A total of 8,812 individuals from two orders (Lepidoptera and Diptera) were observed, including 22 families and 154 species. Bees were the most abundant, with 4,502 individuals, and the most species-rich taxa, with 70 species in five families, followed by flies having 2,509 individuals and 51species in 10 families. Wasps were the least abundant with 1,801 individuals and 33 species in seven families. The assemblage structure of pollinator communities as visualized through rank abundance curves showed that there were many species with low abundance and only a few species with a much higher abundance. The most abundant species among the bees, in order, were Nomia sp.3, Megachile bicolor, and Colletes sp.3; among flies, Syrphus sp.2, Calliphoridae sp.1, and Empididae sp.4; and among wasps, Tiphiidae sp.1, Myzininae sp.2, and Scelionidae sp.1.     

Making learning easy: the acquisition of visual information during the orientation flights of social wasps

The flights of individual wasps (Vespula) were recorded as they approached a small feeder on the ground that was marked by a black cylinder ca 15 cm away. Two navigational strategies are used in these approaches. Initially, the wasp aims at the cylinder, treating it as a beacon and fixating it with frontal retina. In the last stage of the flight, the wasp assumes a preferred orientation so that the cylinder takes up a constant, more peripheral retinal position as the wasp nears the feeder. Path guidance by image-matching is likely to be limited to this final segment of the return. Wasps could gain the information needed for these distinct navigational strategies during the learning flights that they perform on their initial departures from the feeder. They fly away from the feeder in a series of arcs while turning at a mean angular velocity of 226°/s. The cylinder tends to be viewed with frontal retina during the arcs suggesting that the information required for aiming at the cylinder is acquired then. For image matching, the appearance of the cylinder needs to be learnt when the wasp is in the orientation that it adopts close to the feeder on its return flight. Wasps tend to assume this orientation during learning flights while they face the feeder. Such inspections of the feeder occur at the ends of arcs when a wasp's turning velocity is low.     

View-based navigation in Hymenoptera: multiple strategies of landmark guidance in the approach to a feeder

In order to analyse how landmarks guide the last stages of an insect's approach to a goal, we recorded many flights of individual wasps and honeybees as they flew to an inconspicuous feeder on the ground that was marked by one or by two nearby landmarks. An individual tends to approach the feeder from a constant direction, flying close to the ground. Its body is oriented in roughly the same horizontal direction during the approach so that the feeder and landmarks are viewed over a narrow range of directions. Consequently, when the insect arrives at the feeder, the landmarks take up a standard position on the retina. Three navigational strategies govern the final approach. The insect first aims at a landmark, treating it as a beacon. Secondly, bees learn the appearance of a landmark with frontal retina and they associate with this stored view a motor trajectory which brings them from the landmark sufficiently close to the goal that it can be reached by image matching. Insects then move so as to put the landmark in its standard retinal position. Image matching is shown to be accomplished by a control system which has as set points the standard retinal position of the landmark and some parameter related to its retinal size. Accepted: 1 March 1997