Asymmetrical nature of the Trollius-Chiastocheta interaction: insights into the evolution of nursery pollination systems

The mutualistic versus antagonistic nature of an interaction is defined by costs and benefits of each partner, which may vary depending on the environment. Contrasting with this dynamic view, several pollination interactions are considered as strictly obligate and mutualistic. Here, we focus on the interaction between Trollius europaeus and Chiastocheta flies, considered as a specialized and obligate nursery pollination system - the flies are thought to be exclusive pollinators of the plant and their larvae develop only in T.europaeus fruits. In this system, features such as the globelike flower shape are claimed to have evolved in a coevolutionary context. We examine the specificity of this pollination system and measure traits related to offspring fitness in isolated T.europaeus populations, in some of which Chiastocheta flies have gone extinct. We hypothesize that if this interaction is specific and obligate, the plant should experience dramatic drop in its relative fitness in the absence of Chiastocheta. Contrasting with this hypothesis, T.europaeus populations without flies demonstrate a similar relative fitness to those with the flies present, contradicting the putative obligatory nature of this pollination system. It also agrees with our observation that many other insects also visit and carry pollen among T.europaeus flowers. We propose that the interaction could have evolved through maximization of by-product benefits of the Chiastocheta visits, through the male flower function, and selection on floral traits by the most effective pollinator. We argue this mechanism is also central in the evolution of other nursery pollination systems.

Preliminary study of insect attraction by a mixture of semiochemicals containing 1,2,4-Trimethoxybenzene in domestic citric-culture

In this work we describe a new efficient strategy for the preparation of 1,2,4-trimethoxybenzene (3) in 56% overall yield. The compound 3 was used in a preliminary study of insect attraction by a mixture of semiochemicals called TIV, composed of indol (1), vanillin (2) and 1,2,4-trimethoxybenzene (3), in eight Mc Phail style traps installed at a domestic orchard of citric-culture, containing 120 trees not infected by plagues in Bom Jesus Farm, located next to a patch of the Atlantic Forest, at Silva Jardim, Rio de Janeiro, Brazil.

Pollination and other biotic interactions in figs of Ficus eximia Schott (Moraceae)

During the period from 1992 to 1997, interactions of several organisms and Ficus eximia figs, a monoecious species, were studied in plants located in Campinas/SP and Londrina/PR (Brazil). Ficus eximia is pollinated by a single fig wasp species, Pegoscapus sp. (Hymenoptera: Agaonidae, Agaoninae), but also visited by other 14 non-pollinating wasps (Agaonidae, Eurytomidae, Torymidae). Mites (Tarsonemidae), nematodes (Diplogasteridae) and fungi which use the body of the pollinating fig wasp to disperse themselves were also observed.

Pollination ecology of Tabebuia aurea (Manso) Benth. & Hook. and T. ochracea (Cham.) Standl. (Bignoniaceae) in Central Brazil cerrado vegetation

The pollination ecology and breeding systems of Tabebuia aurea (Manso) Benth. & Hook., and T. ochracea (Cham.) Standl. were investigated in an area of cerrado vegetation in the Federal District of Brazil. These species occur sympatrically, flower massively and synchronously for a month, during the dry season (July to September). Both have diurnal anthesis, with similar floral structures, a yellow tubular corolla and produce nectar. Fourteen species of bees visited both Tabebuia species, but, only three Centris species and Bombus morio, were considered potential pollinators, because of their high frequency on the flowers and their efficiency in carrying pollen. Tests on the breeding systems of T. aurea and T. ochracea demonstrated that boths species are self-incompatible, with late-acting self-incompatibility. The proportion of fruit set from cross pollination (T. aurea 17.2% and T. ochracea 12.3%) in both species was low considering the great number of flowers displayed. This suggests a lack of maternal resources for fruit-set. The great amount of seeds per fruit (about 92 in T. aurea and 285 in T. ochracea) may represent an investment of maternal resources allocated on higher quality of fertilized ovules.

Reproductive phenology, pollination and seed dispersal syndromes on sandstone outcrop vegetation in the "Chapada Diamantina", northeastern Brazil: population and community analyses

Vegetation on rock outcrops in the "Chapada Diamantina" (soil islands) is often aggregated and surrounded by nude rock surfaces, thus creating natural units with well defined limits. The flowering and fruiting cycles of plants on 58 soil islands at altitudes between 1,100 and 1,140 meters above sea leavel were analyzed at Mãe Inácia Peak (12°27' S and 41°28' W) in the "Chapada Diamantina", Bahia, Brazil. The presence/absence of flowering and fruiting species on each soil island, and their respective cover areas were analyzed at both the population and community levels, and the phenophases of flowering and fruiting were observed during 24 successive months. The analyses of pollination and seed dispersal syndromes indicated that animals are more important in pollination than in seed dispersal (which is predominantly by anemochory and autochory). The flowering and fruiting of plants with animal pollination syndromes were correlated with rainfall and temperature. The flowering season varied during the year according to the pollination syndrome involved: entomophily was predominant from summer through autumn, ornithophily was predominant during winter, and anemophily in the spring. The staggered timing of flowering and fruiting among different species provides a nearly continuous supply of resources for the local fauna.

Pollination biology in Jacaranda copaia (Aubl.) D. Don. (Bignoniaceae) at the "Floresta Nacional do Tapajós", Central Amazon, Brazil

Jacaranda copaia (Aubl.) D. Don is a pioneer tree widespread in the Brazilian Amazon, usually found colonizing forest gaps and altered areas, and the forest fragment edges. This study investigated aspects of the floral biology, breeding system and pollinators of J. copaia trees. Flowering lasts from August to November, during the low rainfall period extending up to four weeks per tree and 3-4 months for the population as a whole, characterizing a cornucopia flowering pattern. The fruit set ends in the beginning of the rainy season, with wind dispersed winged seeds. Fruit set from open pollination was 1.06% (n = 6,932). Hand pollination using self-pollen (n = 2,099) did not set fruits. Cross-pollination resulted in 6.54% fruit set (n = 2,524), representing six times more than the natural pollination rate (1.06%, n = 6,932). Flowers excluded from insect visitation (automatic self-pollination) did not set fruits (n = 5,372). Pollen tube growth down to ovary was detected under fluorescence microcoscopy in cross-pollinated and selfed pistils. The species is an obligate allogamous plant, with late-acting self-incompatibility system. Approximately 40 species of native bees visited the flowers, but the main pollinators were medium-sized solitary bees as Euglossa and Centris species due to the compatibility between their body sizes with the corolla tube, direct contact with the reproductive structures and high frequency of visits.

The bees of Niagara: a test of the intermediate disturbance hypothesis /

This study examined the bee fauna of the Carolinian Zone in Ontario, Canada. In 2003, 15687 individuals from 152 species of bees were collected. Tliere were many rare species but few abundant species. There were three distinct bee seasons. The Niagara bee assemblage was less diverse compared to other Carolinian Zone assemblages and types of landscapes. This study also examined how anthropogenic disturbance affects the diversity of bee assemblages. The intermediate disturbance hypothesis (IDH) was tested by selecting field sites subject to low, intermediate, and high disturbance. Intermediate disturbance had the highest species richness (SR=1 15) and most bees (N=556I), followed by low disturbance (SR= 100, N=2975), then high disturbance (SR=72, N=1364), supporting the IDH. Increased species richness in areas of intermediate disturbance was due to higher abundance, possibly because more blooming flowers were found there. Bees were larger in high disturbance areas but smaller in areas of high and intermediate disturbance.

Evolutionary Origin and Maintenance of Sociality in the Small Carpenter Bees

Many arthropods exhibit behaviours precursory to social life, including adult longevity, parental care, nest loyalty and mutual tolerance, yet there are few examples of social behaviour in this phylum. The small carpenter bees, genus Ceratina, provide important insights into the early stages of sociality. I described the biology and social behaviour of five facultatively social species which exhibit all of the preadaptations for successful group living, yet present ecological and behavioural characteristics that seemingly disfavour frequent colony formation. These species are socially polymorphic with both / solitary and social nests collected in sympatry. Social colonies consist of two adult females, one contributing both foraging and reproductive effort and the second which remains at the nest as a passive guard. Cooperative nesting provides no overt reproductive benefits over solitary nesting, although brood survival tends to be greater in social colonies. Three main theories explain cooperation among conspecifics: mutual benefit, kin selection and manipulation. Lifetime reproductive success calculations revealed that mutual benefit does not explain social behaviour in this group as social colonies have lower per capita life time reproductive success than solitary nests. Genetic pedigrees constructed from allozyme data indicate that kin selection might contribute to the maintenance of social nesting -, as social colonies consist of full sisters and thus some indirect fitness benefits are inherently bestowed on subordinate females as a result of remaining to help their dominant sister. These data suggest that the origin of sociality in ceratinines has principal costs and the great ecological success of highly eusociallineages occurred well after social origins. Ecological constraints such as resource limitation, unfavourable weather conditions and parasite pressure have long been considered some of the most important selective pressures for the evolution of sociality. I assessed the fitness consequences of these three ecological factors for reproductive success of solitary and social colonies and found that nest sites were not limiting, and the frequency of social nesting was consistent across brood rearing seasons. Local weather varied between seasons but was not correlated with reproductive success. Severe parasitism resulted in low reproductive success and total nest failure in solitary nests. Social colonies had higher reproductive success and were never extirpated by parasites. I suggest that social nesting represents a form of bet-hedging. The high frequency of solitary nests suggests that this is the optimal strategy when parasite pressure is low. However, social colonies have a selective advantage over solitary nesting females during periods of extreme parasite pressure. Finally, the small carpenter bees are recorded from all continents except Antarctica. I constructed the first molecular phylogeny of ceratinine bees based on four gene regions of selected species covering representatives from all continents and ecological regions. Maximum parsimony and Bayesian Inference tree topology and fossil dating support an African origin followed by an Old World invasion and New World radiation. All known Old World ceratinines form social colonies while New World species are largely solitary; thus geography and phylogenetic inertia are likely predictors of social evolution in this genus. This integrative approach not only describes the behaviour of several previously unknown or little-known Ceratina species, bu~ highlights the fact that this is an important, though previously unrecognized, model for studying evolutionary transitions from solitary to social behaviour.

Bud development, flowering and fruit set of Moringa oleifera Lam. (Horseradish Tree) as affected by various irrigation levels

Moringa oleifera is becoming increasingly popular as an industrial crop due to its multitude of useful attributes as water purifier, nutritional supplement and biofuel feedstock. Given its tolerance to sub-optimal growing conditions, most of the current and anticipated cultivation areas are in medium to low rainfall areas. This study aimed to assess the effect of various irrigation levels on floral initiation, flowering and fruit set. Three treatments namely, a 900 mm (900IT), 600 mm (600IT) and 300 mm (300IT) per annum irrigation treatment were administered through drip irrigation, simulating three total annual rainfall amounts. Individual inflorescences from each treatment were tagged during floral initiation and monitored throughout until fruit set. Flower bud initiation was highest at the 300IT and lowest at the 900IT for two consecutive growing seasons. Fruit set on the other hand, decreased with the decrease in irrigation treatment. Floral abortion, reduced pollen viability as well as moisture stress in the style were contributing factors to the reduction in fruiting/yield observed at the 300IT. Moderate water stress prior to floral initiation could stimulate flower initiation, however, this should be followed by sufficient irrigation to ensure good pollination, fruit set and yield.

Dispersal capacity and diet breadth modify the response of wild bees to habitat loss

Habitat loss poses a major threat to biodiversity, and species-specific extinction risks are inextricably linked to life-history characteristics. This relationship is still poorly documented for many functionally important taxa, and at larger continental scales. With data from five replicated field studies from three countries, we examined how species richness of wild bees varies with habitat patch size. We hypothesized that the form of this relationship is affected by body size, degree of host plant specialization and sociality. Across all species, we found a positive species–area slope (z ¼ 0.19), and species traits modified this relationship. Large-bodied generalists had a lower z value than small generalists. Contrary to predictions, small specialists had similar or slightly lower z value compared with large specialists, and small generalists also tended to be more strongly affected by habitat loss as compared with small specialists. Social bees were negatively affected by habitat loss (z ¼ 0.11) irrespective of body size. We conclude that habitat loss leads to clear shifts in the species composition of wild bee communities.

Organic farming in isolated landscapes does not benefit flower-visiting insects and pollination

Organic farming has often been found to provide benefits for biodiversity, but the benefits can depend on the species considered and characteristics of the surrounding landscape. In an intensively farmed area of Northeast Italy we investigated whether isolated organic farms, in a conventionally farmed landscape, provided local benefits for insect pollinators and pollination services. We quantified the relative effects of local management (i.e. the farm system), landscape management (proportion of surrounding uncultivated land) and interactions between them. We compared six organic and six conventional vine fields. The proportion of surrounding uncultivated land was calculated for each site at radii of 200, 500, 1000 and 2000 m. The organic fields did not differ from the conventional in their floral resources or proportion of surrounding uncultivated land. Data were collected on pollinator abundance and species richness, visitation rates to, and pollination of experimental potted plants. None of these factors were significantly affected by the farming system. The abundance of visits to the potted plants in the conventional fields tended to be negatively affected by the proportion of surrounding uncultivated land. The proportion fruit set, weight of seeds per plant and seed weight in conventional and organic fields were all negatively affected by the proportion of surrounding uncultivated land. In vine fields the impact of the surrounding landscape was stronger than the local management. Enhancement of biodiversity through organic farming should not be assumed to be ubiquitous, as potential benefits may be offset by the crop type, organicmanagement practices and the specific habitat requirements in the surrounding landscape.

Conservation ecology of bees: populations, species and communities

Recent concerns regarding the decline of plant and pollinator species, and the impact on ecosystem functioning, has focused attention on the local and global threats to bee diversity. As evidence for bee declines is now accumulating from over broad taxonomic and geographic scales, we review the role of ecology in bee conservation at the levels of species, populations and communities. Bee populations and communities are typified by considerable spatiotemporal variation; whereby autecological traits, population size and growth rate, and plant-pollinator network architecture all play a role in their vulnerability to extinction. As contemporary insect conservation management is broadly based on species- and habitat-targeted approaches, ecological data will be central to integrating management strategies into a broader, landscape scale of dynamic, interconnected habitats capable of delivering bee conservation in the context of global environmental change.

Linking bees and flowers: How do floral communities structure pollinator communities?

Pollinators provide essential ecosystem services, and declines in some pollinator communities around the world have been reported. Understanding the fundamental components defining these communities is essential if conservation and restoration are to be successful. We examined the structure of plant-pollinator communities in a dynamic Mediterranean landscape, comprising a mosaic of post-fire regenerating habitats, and which is a recognized global hotspot for bee diversity. Each community was characterized by a highly skewed species abundance distribution, with a few dominant and many rare bee species, and was consistent with a log series model indicating that a few environmental factors govern the community. Floral community composition, the quantity and quality of forage resources present, and the geographic locality organized bee communities at various levels: (1) The overall structure of the bee community (116 species), as revealed through ordination, was dependent upon nectar resource diversity (defined as the variety of nectar volume-concentration combinations available), the ratio of pollen to nectar energy, floral diversity, floral abundance, and post-fire age. (2) Bee diversity, measured as species richness, was closely linked to floral diversity (especially of annuals), nectar resource diversity, and post-fire age of the habitat. (3) The abundance of the most common species was primarily related to post-fire age, grazing intensity, and nesting substrate availability. Ordination models based on age-characteristic post-fire floral community structure explained 39-50% of overall variation observed in bee community structure. Cluster analysis showed that all the communities shared a high degree of similarity in their species composition (27-59%); however, the geographical location of sites also contributed a smaller but significant component to bee community structure. We conclude that floral resources act in specific and previously unexplored ways to modulate the diversity of the local geographic species pool, with specific disturbance factors, superimposed upon these patterns, mainly affecting the dominant species.

Social bees and food plant associations in the Nilgiri Biosphere Reserve, India

The diversity of social bees was assessed at 15 sites across five locations of the Nilgiri Biosphere Reserve, Western Ghats, India, from January to December 2007. We also conducted floristic analyses of local vegetation in each site using one-hectare sample plots. All woody species with a dbh (diameter at breast height) : 30 cm were recorded within the plots. A total area of 9.72 ha was assessed for floristic composition. Similarity of floristic composition between sites was determined using the Jaccard's distance measure and a dendrogram constructed based on the hierarchical clustering of floristic dissimilarities between sites. A Bee Importance Index (BII) was developed to give a measure of the bee diversity at each site. This index was a sum of the species richness of bee species in a site and their visitation frequencies to flowers, calculated as mean flower visits hour 1 within 2 focal patches within one hectare plots. The visits of bee species to flowers were also recorded. The Jaccard distance measure indicated that the montane sites were quite dissimilar to the low elevation sites in floristic diversity. The BII was 7-9 for the wet forest sites and ranged from 4-6 for drier forest sites. Seventy three plant species were identified as social bee plants and of them 45% were visited by one species of bee, 37% by two bee species and 18% by more than two bee species, indicating a certain degree of floral specialization among bees.

Local management and landscape drivers of pollination and biological control services in a Kenyan agro-ecosystem

Arthropods that have a direct impact on crop production (i.e. pests, natural enemies and pollinators) can be influenced by both local farm management and the context within which the fields occur in the wider landscape. However, the contributions and spatial scales at which these drivers operate and interact are not fully understood, particularly in the developing world. The impact of both local management and landscape context on insect pollinators and natural enemy communities and on their capacity to deliver related ecosystem services to an economically important tropical crop, pigeonpea was investigated. The study was conducted in nine paired farms across a gradient of increasing distance to semi-native vegetation in Kibwezi, Kenya. Results show that proximity of fields to semi-native habitats negatively affected pollinator and chewing insect abundance. Within fields, pesticide use was a key negative predictor of pollinator, pest and foliar active predator abundance. On the contrary, fertilizer application significantly enhanced pollinator and both chewing and sucking insect pest abundance. At a 1 km spatial scale of fields, there were significant negative effects of the number of semi-native habitat patches within fields dominated by mass flowering pigeonpea on pollinators abundance. For service provision, a significant decline in fruit set when insects were excluded from flowers was recorded. This study reveals the interconnections of pollinators, predators and pests with pigeonpea crop. For sustainable yields and to conserve high densities of both pollinators and predators of pests within pigeonpea landscapes, it is crucial to target the adoption of less disruptive farm management practices such as reducing pesticide and fertilizer inputs.