Density of predating Asian hornets at hives disturbs the 3D flight performance of honey bees and decreases predation success.

Automated 3D image-based tracking systems are new and promising devices to investigate the foraging behavior of flying animals with great accuracy and precision. 3D analyses can provide accurate assessments of flight performance in regard to speed, curvature, and hovering. However, there have been few applications of this technology in ecology, particularly for insects. We used this technology to analyze the behavioral interactions between the Western honey bee Apis mellifera and its invasive predator the Asian hornet, Vespa velutina nigrithorax. We investigated whether predation success could be affected by flight speed, flight curvature, and hovering of the Asian hornet and honey bees in front of one beehive. We recorded a total of 603,259 flight trajectories and 5175 predator-prey flight interactions leading to 126 successful predation events, representing 2.4% predation success. Flight speeds of hornets in front of hive entrances were much lower than that of their bee prey; in contrast to hovering capacity, while curvature range overlapped between the two species. There were large differences in speed, curvature, and hovering between the exit and entrance flights of honey bees. Interestingly, we found hornet density affected flight performance of both honey bees and hornets. Higher hornet density led to a decrease in the speed of honey bees leaving the hive, and an increase in the speed of honey bees entering the hive, together with more curved flight trajectories. These effects suggest some predator avoidance behavior by the bees. Higher honey bee flight curvature resulted in lower hornet predation success. Results showed an increase in predation success when hornet number increased up to 8 individuals, above which predation success decreased, likely due to competition among predators. Although based on a single colony, this study reveals interesting outcomes derived from the use of automated 3D tracking to derive accurate measures of individual behavior and behavioral interactions among flying species.

Quantifying the impact of an invasive Hornet on Bombus terrestris Colonies.

The invasive hornet Vespa velutina nigrithorax is considered a proliferating threat to pollinators in Europe and Asia. While the impact of this species on managed honey bees is well-documented, effects upon other pollinator populations remain poorly understood. Nonetheless, dietary analyses indicate that the hornets consume a diversity of prey, fuelling concerns for at-risk taxa. Here, we quantify the impact of V. velutina upon standardised commercially-reared colonies of the European bumblebee, Bombus terrestris terrestris. Using a landscape-scale experimental design, we deploy colonies across a gradient of local V. velutina densities, utilising automated tracking to non-invasively observe bee and hornet behaviour, and quantify subsequent effects upon colony outcomes. Our results demonstrate that hornets frequently hunt at B. terrestris colonies, being preferentially attracted to those with high foraging traffic, and engaging in repeated-yet entirely unsuccessful-predation attempts at nest entrances. Notably however, we show that B. terrestris colony weights are negatively associated with local V. velutina densities, indicating potential indirect effects upon colony growth. Taken together, these findings provide the first empirical insight into impacts on bumblebees at the colony level, and inform future mitigation efforts for wild and managed pollinators.

Hornets and Honey Bees: A Coevolutionary Arms Race between Ancient Adaptations and New Invasive Threats.

Hornets and honey bees have a long history of coevolution resulting in a plethora of captivating adaptations and counteradaptations between predator and prey. From simple physiological mechanisms to complex behavioral strategies, some Vespa hornets have specialized in hunting honey bees, while the latter have put in place effective defenses to counteract their attack. Both hornets and honey bees have evolved the ability to detect the odors and the pheromones emitted by the other to locate the prey or to spot foraging predators. Hornets often rely on their bigger size, heavily armored body and destructive attacks, while honey bees differentiated collective defense responses finely coordinated to deter or kill the hornet menace. However, when new species of hornets and honey bees come into contact, the absence of coevolution can have a heavy impact on the defenseless bees. The evolutionary arms race between hornets and honey bees provides not only compelling examples of adaptations and counteradaptations between predator and prey, but could also represent a starting point for the development of effective and sustainable strategies to protect honey bees and beekeeping activities and to control invasive alien species of hornets.

Comparison of reproductive traits of foundresses in a native and an invasive hornet in Europe.

The population dynamics of annual social hymenoptera such as vespids depend largely on the fertility of the foundresses, which, in turn, is a key factor in the context of biological invasions. The native European hornet Vespa crabro (Vc) and the invasive Asian hornet Vespa velutina (Vv) have generally similar ecological traits, e.g. nesting and feeding habits, although they differ in colony size, which is higher in Vv. Furthermore, in contrast to Vc, Vv is more specialized in its predatory habits, intensively hunting honey bees at the hive. Comparing the morphological and reproductive traits of two closely related species occupying the same ecological niche, one of which is a native species and the other an alien, can help us to gain an understanding of the invasion process. To this end, we here compare reproductive (ovarian size and maturation, fat level, spermatheca size and sperm stock, fecundity) and morphological traits (head size, weight) of the foundresses of these two hornet species. We observed that ovarian maturation began approximately one month earlier in Vv than in Vc, and that the fat level in the former was lower. We found twice the number of sperm in the mated foundresses of Vv than in those of Vc (more than 100 × 103 and less than 50 × 103 sperm, respectively), in a 16% smaller spermatheca in Vc. Furthermore, the sperm of Vv was found to be 65% shorter than that of Vc. The precocity and higher potential fecundity of Vv queens may have favoured this species over Vc in terms of predatory behaviour, and thereby contributed to its invasiveness.

Activity rhythm and action range of workers of the invasive hornet predator of honeybees Vespa velutina, measured by radio frequency identification tags.

In social insects, the activity rhythm of foragers and their action range determinate the activity of the colony. In vespids, which are mostly predators, the foraging range of workers determines their maximum predation pressure round the nest. One of these species, Vespa velutina, a recently invasive species introduced into Europe, exerts a strong predation on honeybees at the hive. Therefore, the definition of its activity rhythm and spatial range of predation is of primary importance. Using radio frequency identification tags (RFID), two experiments were carried out to (a) determine their return ability (called homing) in releasing 318 individuals at different distance from their colony and (b) monitor their foraging activity rhythm and the duration of their flights based on 71 individuals followed 24 hr/24 during 2 months. The homing ability of V. velutina was evaluated to be up to 5,000 m and was not affected by the cardinal orientation of release point. The lag time to return to the nest increased with the distance of release. Most of the flight activity took place between 07:00 a.m. and 08:00 p.m., hornets doing principally short flights of less than an hour. Foraging range was thus estimated ca. 1,000 m around the nest. This study of V. velutina assisted by RFID tags provides for the first time a baseline for its potential foraging distance that increase our knowledge of this species to (a) refine more accurately models for risk assessment and (b) define security perimeter for early detection of predation on invasion front.

Delayed sexual maturity in males of Vespa velutina.

Vespa velutina var nigrithorax (Lepelletier, 1835) is an invasive predator of bees accidentally introduced in France in 2004, and it is having a serious impact on apiculture and ecosystems. Studying the reproduction of an invasive species is key to assess its population dynamic. This study explores the sexual maturation of V. velutina males and the evolution of their fertility. The main studied parameters were physiologic (spermiogenesis, spermatogenesis) and anatomic (testes size and structure, head width). Two populations of males were described based on their emergence period: early males in early summer or classic males in autumn. Each testis has an average of 108 testicular follicles. Spermatogenesis is synchronous, with only 1 sperm production wave, and completed, on average, at 10.3 d after emergence with the degeneration of the testes. The sperm counts in seminal vesicles of mature males are 3 × 106 in October/November and 0.8 × 106 in June. In comparison, females store 0.1 × 106 sperm in their spermathecae. The early males emerged from colonies made by fertilized queens. The reproductive potential of these early males seemed limited, and their function in the colony is discussed. The sperm stock evolution in autumn males suggests the occurrence of a reproductive pattern of male competition for the access to females and a single copulation per male. The synchronicity of male and foundress emergences and sexual maturation is of primary importance for the mating success and the future colony development.

Searching for nests of the invasive Asian hornet (Vespa velutina) using radio-telemetry.

Asian hornets (Vespa velutina) are voracious predators of bees, and are the latest emerging threat to managed and wild pollinator populations in Europe. To prevent establishment or reduce the rate of spread of V. velutina, early detection and destruction of nests is considered the only option. Detection is difficult as their nests are well hidden and flying hornets are difficult to follow over long distances. We address this challenge by tracking individual V. velutina workers flying back to their nests using radio telemetry for the first time, finding five previously undiscovered nests, up to 1.33 km from hornet release points. Hornets can fly with 0.28 g tags if the tag:hornet ratio is less than 0.8. This method offers a step-change in options to tackle the spread of this invader, providing an efficient means of finding V. velutina nests in complex environments to manage this emerging threat to pollinators.

Behavioral syndrome in a native and an invasive hymenoptera species.

Recent studies have focused on the role of behavior in biological invasions. Individuals may differ consistently in time for several behavioral traits (personality) which covary (behavioral syndrome) resulting in different behavioral types, some of them favoring invasion. Social hymenopterans have a strong potential to be invaders and their success depends primarily on the foundresses' ability to found viable colonies. They are expected to be active, explorative and bold for optimally establishing their nest. In Europe, 2 hornet species coexist: the native Vespa crabro and the invasive Vespa velutina. These 2 species may compete for nesting sites and we suggest that the initial success of V. velutina has been favored by its behavior in outperforming V. crabro for the traits involved in nest initiation. Here, we (i) defined the personality of V. crabro and V. velutina, (ii) tested for the existence of behavioral syndrome in these species, and (iii) compared their performances using an open-field test. Our results show that V. crabro foundresses behave consistently but not V. velutina; this lack of consistency being mainly due to reduced variance among individuals. This result questions the possibility of detecting consistent behavioral differences in species having recently undergone a strong bottleneck. Both species exhibit the same correlations between activity, boldness and exploration and V. velutina clearly outperforms V. crabro for all traits. Our results suggest that activity, boldness, and exploration are implicated in both hornet nest initiation and invasion process which contributed to explain why social hymenopterans are so successful at colonization.