Checklist of the bees (Hymenoptera, Apoidea) of New Caledonia.

Background: In a world where insects and notably bees are declining, assessing their distribution over time and space is crucial to evaluate species status and highlight conservation priorities. However, this can be a daunting task, especially in areas such as tropical oceanic islands where exhaustive samplings over time have been lacking. This is the case in New Caledonia, an archipelago located in the southwest Pacific. Historical records of bee species are piecemeal and, although contemporary samplings have significantly advanced our knowledge of the bee fauna of New Caledonia, the status of several species remains to be elucidated. New information: Here, we provide an updated checklist of the 51 bee species recorded for New Caledonia using previous publications and personal samplings. We documented their distribution, origin (i.e. endemic, native or alien) and the year and location of their occurrences. Based on the year of their first capture and the year of their last capture, we determined an occurrence status for each species. Thus, 10 years after the last checklist of the New Caledonian bee fauna, the literature review and recent samplings allowed us to add six new species to the list. Half of them are recently introduced species including one firstly mentioned in this paper (i.e. Hylaeusalbonitens). We consider here that 30 species are effectively present on the territory and the presence of 21 species could not be determined due to a lack of data, which highlights the need to increase sampling efforts across New Caledonia. Given the difficulty of exhaustively sampling the entire archipelago, we would recommend taking, as a starting point, altitude environments and areas where data-deficient species were captured. In a broader perspective, biomolecular analyses are crucial to confirm species identifications. This is also needed to make comparisons between archipelagoes and thus clarify the distribution and status of species at the scale of the southwest Pacific.

Effects of litter quality on foraging behaviour and demographic parameters in Folsomia candida (Collembola).

Litter quality has long been associated with demographic parameters of Collembola populations. However, little is known about the capacity of Collembola to perceive and seek better litter quality. To address this gap, three complementary laboratory experiments were carried out with the Collembola Folsomia candida. First, populations were fed on three different types of leaf litters (Quercus pubescens, Acer opalus and Prunus avium) and a control (agar-agar-brewer's yeast mixture) for 6 weeks to assess their impacts on demography (reproduction rate and population size). Second, the body length of individuals differentially fed with the same four types of resources was measured to assess a functional trait that can potentially affect movement parameters such as prospected area or foraging speed. Third, F. candida single individuals were exposed to the same litter quality gradient and placed at an increasing distance from the litter (from 1 to 5 cm). For 10 min, their foraging behaviour was recorded which included prospected area, foraging speed, perception distance and success in reaching the litter (foraging success). As expected, low-quality litter (i.e. Q. pubescens) contributed to low population growth compared to the control treatment and the high-quality litters (P. avium and A. opalus). In the third experiment, the probability of finding the resource was negatively correlated to the distance, but was unrelated to the litter quality and the Collembola body length. When resource was perceived, F. candida was able to switch from non-directional to directional movements, with a large variability in the perception distance from a few millimetres to several centimetres. Taken together, our results indicate that litter quality plays a relevant role in Collembola demographic parameters once the population settles on litter patch, but not on foraging behaviour to select high-quality resources.

Microbiota and pathogens in an invasive bee: Megachile sculpturalis from native and invaded regions.

The present study aimed to characterise the bacterial, fungal and parasite gut community of the invasive bee Megachile sculpturalis sampled from native (Japan) and invaded (USA and France) regions via 16S rRNA and ITS2 amplicon sequencing and PCR detection of bee microparasites. The bacterial and fungal gut microbiota communities in bees from invaded regions were highly similar and differed strongly from those obtained in Japan. Core amplicon sequence variants (ASVs) within each population represented environmental micro-organisms commonly present in bee-associated niches that likely provide beneficial functions to their host. Although the overall bacterial and fungal communities of the invasive M. sculpturalis in France and the co-foraging native bees Anthidium florentinum and Halictus scabiosae, were significantly different, five out of eight core ASVs were shared suggesting common environmental sources and potential transmission. None of the 46 M. sculpturalis bees analysed harboured known bee pathogens, while microparasite infections were common in A. florentinum, and rare in H. scabiosae. A common shift in the gut microbiota of M. sculpturalis in invaded regions as a response to changed environmental conditions, or a founder effect coupled to population re-establishment in the invaded regions may explain the observed microbial community profiles and the absence of parasites. While the role of pathogen pressure in shaping biological invasions is still debated, the absence of natural enemies may contribute to the invasion success of M. sculpturalis.

Apoidea of the collections of Lyon, Aix-en-Provence, Marseille and Toulon Museums of Natural History (France).

Background: Many insect species have shown dramatic declines over the last decades, as a result of man-related environmental changes. Many species which were formerly widespread are now rare. To document this trend with evidence, old records of collected specimens are vital. New information: We provide here the data on 9752 bee (Hymenoptera: Apoidea) specimens hosted in several museums of south-east France: Musée des Confluences in Lyon, Muséum d'Histoire Naturelle de Marseille, Muséum d'Aix-en-Provence and the Muséum Départemental du Var in Toulon. Most of the specimens (9256) come from France and include data on 552 named species. For most of these specimens, the geographical location, including geographical coordinates, is based on the locality (town or village) where they were collected. The specimens were captured from the beginning of the nineteenth century to 2018. The identifications of 1377 specimens, mainly belonging to the genus Bombus, are considered reliable, as these were performed or been checked since 2009. All the other reported identifications are the original ones given by the original collectors.

A large-scale dataset reveals taxonomic and functional specificities of wild bee communities in urban habitats of Western Europe.

Wild bees are declining, mainly due to the expansion of urban habitats that have led to land-use changes. Effects of urbanization on wild bee communities are still unclear, as shown by contrasting reports on their species and functional diversities in urban habitats. To address this current controversy, we built a large dataset, merging 16 surveys carried out in 3 countries of Western Europe during the past decades, and tested whether urbanization influences local wild bee taxonomic and functional community composition. These surveys encompassed a range of urbanization levels, that were quantified using two complementary metrics: the proportion of impervious surfaces and the human population density. Urban expansion, when measured as a proportion of impervious surfaces, but not as human population density, was significantly and negatively correlated with wild bee community species richness. Taxonomic dissimilarity of the bee community was independent of both urbanization metrics. However, occurrence rates of functional traits revealed significant differences between lightly and highly urbanized communities, for both urbanization metrics. With higher human population density, probabilities of occurrence of above-ground nesters, generalist and small species increased. With higher soil sealing, probabilities of occurrence of above-ground nesters, generalists and social bees increased as well. Overall, these results, based on a large European dataset, suggest that urbanization can have negative impacts on wild bee diversity. They further identify some traits favored in urban environments, showing that several wild bee species can thrive in cities.

On the road: Anthropogenic factors drive the invasion risk of a wild solitary bee species.

Complex biotic networks of invaders and their new environments pose immense challenges for researchers aiming to predict current and future occupancy of introduced species. This might be especially true for invasive bees, as they enter novel trophic interactions. Little attention has been paid to solitary, invasive wild bees, despite their increasing recognition as a potential global threat to biodiversity. Here, we present the first comprehensive species distribution modelling approach targeting the invasive bee Megachile sculpturalis, which is currently undergoing parallel range expansion in North America and Europe. While the species has largely colonised the most highly suitable areas of North America over the past decades, its invasion of Europe seems to be in its early stages. We showed that its current distribution is largely explained by anthropogenic factors, suggesting that its spread is facilitated by road and maritime traffic, largely beyond its intrinsic dispersal ability. Our results suggest that M. sculpturalis is likely to be negatively affected by future climate change in North America, while in Europe the potential suitable areas at-risk of invasion remain equally large. Based on our study, we emphasise the role of expert knowledge for evaluation of ecologically meaningful variables implemented and interpreted for species distribution modelling. We strongly recommend that the monitoring of this and other invasive pollinator species should be prioritised in areas identified as at-risk, alongside development of effective management strategies.

Landscape and Local Drivers Affecting Flying Insects along Fennel Crops (Foeniculum vulgare, Apiaceae) and Implications for Its Yield.

Agricultural landscapes are increasingly characterized by intensification and habitat losses. Landscape composition and configuration are known to mediate insect abundance and richness. In the context of global insect decline, and despite 75% of crops being dependent on insects, there is still a gap of knowledge about the link between pollinators and aromatic crops. Fennel (Foeniculum vulgare) is an aromatic plant cultivated in the South of France for its essential oil, which is of great economic interest. Using pan-traps, we investigated the influence of the surrounding habitats at landscape scale (semi-natural habitat proportion and vicinity, landscape configuration) and local scale agricultural practices (insecticides and patch size) on fennel-flower-visitor abundance and richness, and their subsequent impact on fennel essential oil yield. We found that fennel may to be a generalist plant species. We did not find any effect of intense local management practices on insect abundance and richness. Landscape configuration and proximity to semi-natural habitat were the main drivers of flying insect family richness. This richness positively influenced fennel essential oil yield. Maintaining a complex configuration of patches at the landscape scale is important to sustain insect diversity and crop yield.

Wild insect diversity increases inter-annual stability in global crop pollinator communities.

While an increasing number of studies indicate that the range, diversity and abundance of many wild pollinators has declined, the global area of pollinator-dependent crops has significantly increased over the last few decades. Crop pollination studies to date have mainly focused on either identifying different guilds pollinating various crops, or on factors driving spatial changes and turnover observed in these communities. The mechanisms driving temporal stability for ecosystem functioning and services, however, remain poorly understood. Our study quantifies temporal variability observed in crop pollinators in 21 different crops across multiple years at a global scale. Using data from 43 studies from six continents, we show that (i) higher pollinator diversity confers greater inter-annual stability in pollinator communities, (ii) temporal variation observed in pollinator abundance is primarily driven by the three-most dominant species, and (iii) crops in tropical regions demonstrate higher inter-annual variability in pollinator species richness than crops in temperate regions. We highlight the importance of recognizing wild pollinator diversity in agricultural landscapes to stabilize pollinator persistence across years to protect both biodiversity and crop pollination services. Short-term agricultural management practices aimed at dominant species for stabilizing pollination services need to be considered alongside longer term conservation goals focussed on maintaining and facilitating biodiversity to confer ecological stability.

Evidence for multiple introductions of an invasive wild bee species currently under rapid range expansion in Europe.

BACKGROUND: Invasive species are increasingly driving biodiversity decline, and knowledge of colonization dynamics, including both drivers and dispersal modes, are important to prevent future invasions. The bee species Megachile sculpturalis (Hymenoptera: Megachilidae), native to East-Asia, was first recognized in Southeast-France in 2008, and has since spread throughout much of Europe. The spread is very fast, and colonization may result from multiple fronts. RESULT: To track the history of this invasion, codominant markers were genotyped using Illumina sequencing and the invasion history and degree of connectivity between populations across the European invasion axis were investigated. Distinctive genetic clusters were detected with east-west differentiations in Middle-Europe. CONCLUSION: We hypothesize that the observed cluster formation resulted from multiple, independent introductions of the species to the European continent. This study draws a first picture of an early invasion stage of this wild bee and forms a foundation for further investigations, including studies of the species in their native Asian range and in the invaded range in North America.

Effects of Urbanization on Plant-Pollinator Interactions in the Tropics: An Experimental Approach Using Exotic Plants.

Land-use changes through urbanization and biological invasions both threaten plant-pollinator networks. Urban areas host modified bee communities and are characterized by high proportions of exotic plants. Exotic species, either animals or plants, may compete with native species and disrupt plant-pollinator interactions. These threats are heightened in insular systems of the Southwest Pacific, where the bee fauna is generally poor and ecological networks are simplified. However, the impacts of these factors have seldom been studied in tropical contexts. To explore those questions, we installed experimental exotic plant communities in urban and natural contexts in New Caledonia, a plant diversity hotspot. For four weeks, we observed plant-pollinator interactions between local pollinators and our experimental exotic plant communities. We found a significantly higher foraging activity of exotic wild bees within the city, together with a strong plant-pollinator association between two exotic species. However, contrary to our expectations, the landscape context (urban vs. natural) had no effect on the activity of native bees. These results raise issues concerning how species introduced in plant-pollinator networks will impact the reproductive success of both native and exotic plants. Furthermore, the urban system could act as a springboard for alien species to disperse in natural systems and even invade them, leading to conservation concerns.

Below-ground competition alters attractiveness of an insect-pollinated plant to pollinators.

Competitive interactions between plants can affect patterns of allocation to reproductive structures through modulation of resource availability. As floral traits involved in plant attractiveness to pollinators can be sensitive to these resources, competition with any neighbouring species may influence the attractiveness of insect-pollinated plants. While pollination research has primarily focused on above-ground interactions, this study aims at investigating if the presence of a competitor plant can modulate neighbouring insect-pollinated plant attractiveness to pollinators and resulting fecundity, especially through below-ground competitive interactions for soil resources. We set up a plot experiment in which we grew an insect-pollinated plant, Sinapis alba (Brassicaceae), in a mixture dominated by a wind-pollinated plant, Holcus lanatus (Poaceae). Individuals of S. alba were either subjected to or isolated from (with buried tubes in the soil) below-ground competition. Across the flowering season, floral traits involved in attractiveness of S. alba and pollinator visitation were followed at the plot and plant level to investigate different scales of attractiveness. At the end of the experiment, seeds were harvested to assess plant fecundity. Competition had a significant negative effect on plot and plant floral display size as well as flower size while nectar traits were not affected. When plants of S. alba were in competition, the time to first visit was altered: the proportion of plots that received a visit was smaller for a given time; in other words, it took more time for a given proportion of plots to be visited and some plots were even never visited. Moreover, pollinators made fewer visits per plots. The proportion of viable seeds produced by S. alba in competition was lower and probably linked to the competition itself rather than changes in pollinator visitation. This study suggests that competitive interactions between plants can modulate pollination interactions even when competing plant species are not insect-pollinated.

Pollinator Specific Richness and Their Interactions With Local Plant Species: 10 Years of Sampling in Mediterranean Habitats.

In the context of global pollinator decline, little is known about the protection status and ecology of many species. This lack of knowledge is particularly important for Mediterranean protected areas that harbor diverse pollinator communities and are subject to considerable anthropogenic pressures. Calanques National Park (85 km2), which is located near Marseille (France), is dominated by Mediterranean low-vegetation habitats, such as phrygana and scrublands. These habitats offer favorable conditions for pollinator species due to the important amount of floral resources. Within a 10-yr period, we recorded bee (Hymenoptera: Apoidea: Anthophila), hover fly (Diptera: Syrphidae), and bee fly (Diptera: Bombyliidae) species and their interactions with the local flora through 10 field campaigns. We caught 250 pollinator species, including 192 bees, 38 hover flies, and 20 bee flies, for a total of 2,770 specimens. We recorded seven threatened bees (six near threatened and one endangered). Among the bee species, 47.9% were below-ground nesting species, and 54.7% were generalist species. Analysis of the pollination network showed that generalist and specialist pollinators do not share the same floral resources. The Cistaceae plant family (Malvales: Cistaceae) acted as a central node in the plant-pollinator network, interacting with 52 different pollinator species, which shows the importance of large open flowers that could be easily visited by both short and long-tongued pollinators in Mediterranean habitats. The occurrence of pollinator species and their ecological traits should strongly contribute to reinforcing the available information to provide or ameliorate the conservation statuses determined by IUCN Red List.

Wild pollinator activity negatively related to honey bee colony densities in urban context.

As pollinator decline is increasingly reported in natural and agricultural environments, cities are perceived as shelters for pollinators because of low pesticide exposure and high floral diversity throughout the year. This has led to the development of environmental policies supporting pollinators in urban areas. However, policies are often restricted to the promotion of honey bee colony installations, which resulted in a strong increase in apiary numbers in cities. Recently, competition for floral resources between wild pollinators and honey bees has been highlighted in semi-natural contexts, but whether urban beekeeping could impact wild pollinators remains unknown. Here, we show that in the city of Paris (France), wild pollinator visitation rates are negatively correlated to honey bee colony densities present in the surrounding landscape (500m -slope = -0.614; p = 0.001 -and 1000m -slope = -0.489; p = 0.005). Regarding the morphological groups of wild pollinators, large solitary bee and beetle visitation rates were negatively affected by honey bee colony densities within a 500m buffer (slope = -0.425, p = 0.007 and slope = - 0.671, p = 0.002, respectively) and bumblebee visitation rates were negatively affected by honey bee colony density within a 1000m buffer (slope = - 0.451, p = 0.012). Further, lower interaction evenness in plant-pollinator networks was observed with high honey bee colony density within a 1000m buffer (slope = -0.487, p = 0.008). Finally, honey bees tended to focus their foraging activity on managed rather than wild plant species (student t-test, p = 0.001) whereas wild pollinators equally visited managed and wild species. We advocate responsible practices mitigating the introduction of high density of honey bee colonies in urban environments. Further studies are however needed to deepen our knowledge about the potential negative interactions between wild and domesticated pollinators.

Range expansion of the Asian native giant resin bee Megachile sculpturalis (Hymenoptera, Apoidea, Megachilidae) in France.

In 2008, a new species for the French bee fauna was recorded in Allauch near Marseille: the giant resin bee, Megachile sculpturalis (Smith, 1853). This was the first European record of this species that is native to East Asia. To our knowledge, it is the first introduced bee species in Europe. Here, we provide an overview of the current distribution of M. sculpturalis in France and we describe the history of its range expansion. Besides our own observations, information was compiled from literature and Internet websites, and by contacting naturalist networks. We collected a total of 117 records (locality × year combinations) for the 2008-2016 period. The geographical range of M. sculpturalis has extended remarkably, now occupying a third of continental France, with the most northern and western records located 335 and 520 km from Allauch, respectively. Information on its phenology, feeding, and nesting behavior is also provided. We report several events of nest occupation or eviction of Osmia sp. and Xylocopa sp. individuals by M. sculpturalis. Our results show that M. sculpturalis is now well established in France. Given its capacity to adapt and rapidly expand its range, we recommend amplifying the monitoring of this species to better anticipate the changes in its geographical range and its potential impacts on native bees.

The proportion of impervious surfaces at the landscape scale structures wild bee assemblages in a densely populated region.

Given the predicted expansion of cities throughout the world, understanding the effect of urbanization on bee fauna is a major issue for the conservation of bees. The aim of this study was to understand how urbanization affects wild bee assemblages along a gradient of impervious surfaces and to determine the influence of landscape composition and floral resource availability on these assemblages. We chose 12 sites with a proportion of impervious surfaces (soil covered by parking, roads, and buildings) ranging from 0.06% to 64.31% within a 500 m radius. We collected using pan trapping and estimated the landscape composition of the sites within a 500 m radius and the species richness of plant assemblages within a 200 m radius. We collected 1104 bees from 74 species. The proportion of impervious surfaces at the landscape scale had a negative effect on wild bee abundance and species richness, whereas local flower composition had no effect. Ground-nesting bees were particularly sensitive to the urbanization gradient. This study provides new evidences of the impact of urbanization on bee assemblages and the proportion of impervious surfaces at the landscape scale emerged as a key factor that drives those assemblages.

Plant pollinator networks along a gradient of urbanisation.

BACKGROUND: Habitat loss is one of the principal causes of the current pollinator decline. With agricultural intensification, increasing urbanisation is among the main drivers of habitat loss. Consequently studies focusing on pollinator community structure along urbanisation gradients have increased in recent years. However, few studies have investigated how urbanisation affects plant-pollinator interaction networks. Here we assessed modifications of plant-pollinator interactions along an urbanisation gradient based on the study of their morphological relationships. METHODOLOGY/PRINCIPAL FINDINGS: Along an urbanisation gradient comprising four types of landscape contexts (semi-natural, agricultural, suburban, urban), we set up experimental plant communities containing two plant functional groups differing in their morphological traits ("open flowers" and "tubular flowers"). Insect visitations on these communities were recorded to build plant-pollinator networks. A total of 17 857 interactions were recorded between experimental plant communities and flower-visitors. The number of interactions performed by flower-visitors was significantly lower in urban landscape context than in semi-natural and agricultural ones. In particular, insects such as Syrphidae and solitary bees that mostly visited the open flower functional group were significantly impacted by urbanisation, which was not the case for bumblebees. Urbanisation also impacted the generalism of flower-visitors and we detected higher interaction evenness in urban landscape context than in agricultural and suburban ones. Finally, in urban context, these modifications lowered the potential reproductive success of the open flowers functional group. CONCLUSIONS/SIGNIFICANCE: Our findings show that open flower plant species and their specific flower-visitors are especially sensitive to increasing urbanisation. These results provide new clues to improve conservation measures within urbanised areas in favour of specialist flower-visitors. To complete this functional approach, studies using networks resolved to the species level along urbanised gradients would be required.