Multiple mating by both sexes in an invasive insect species, Aethina tumida (Coleoptera: Nitidulidae).

Multiple mating by both sexes is common among sexually reproducing animals. Small hive beetles (SHB), Aethina tumida, are parasites of bee nests endemic to sub-Saharan Africa and have become a widespread invasive species. Despite the considerable economic damages they can cause, their basic biology remains poorly understood. Here we show that male and female small hive beetles can mate multiple times, suggesting that costs for mating are low in this species. In an invasive A. tumida population in the United States, a combination of laboratory experiments for males and paternity analysis with eight polymorphic DNA microsatellite markers for field-caught females were used to estimate the number of mating by both sexes. The data show that females and males can mate multiple times-females mated with up to eight males, whereas males mated with at least seven females. The results also showed that A. tumida displayed a skewed paternity, although this was not consistent among the tested females. Thus, first or last male advantage seem to be unlikely in A. tumida. Our observations that individuals of both sexes of A. tumida can mate multiple times opens new research avenues for examining drivers of multiple mating and determining the role it may play in promoting biological invasions.

Cuticular Hydrocarbon Profile of Parasitic Beetles, Aethina tumida (Coleoptera: Nitidulidae).

Cuticular hydrocarbons (CHCs) cover insects' bodies and play important roles in chemical communication, including nestmate recognition, for social insects. To enter colonies of a social host species, parasites may acquire host-specific CHCs or covertly maintain their own CHC profile by lowering its quantity. However, the chemical profile of small hive beetles (SHBs), Aethina tumida, which are parasites of honey bee, Apis mellifera, colonies, and other bee nests, is currently unknown. Here, adults of SHB and honey bee host workers were collected from the same field colonies and their CHC profiles were analysed using GC-MS. The chemical profiles of field-sampled SHBs were also compared with those of host-naive beetles reared in the laboratory. Laboratory-reared SHBs differed in their CHC profiles from field-sampled ones, which showed a more similar, but ten-fold lower, generic host CHC profile compared to host workers. While the data confirm colony-specific CHCs of honey bee workers, the profile of field-collected SHBs was not colony-specific. Adult SHBs often commute between different host colonies, thereby possibly preventing the acquisition of a colony-specific CHC profiles. An ester was exclusive to both groups of SHBs and might constitute an intraspecific recognition cue. Our data suggest that SHBs do not use any finely tuned chemical strategy to conceal their presence inside host colonies and instead probably rely on their hard exoskeleton and defence behaviours.

Evolution of starvation resistance in an invasive insect species, Aethina tumida (Coleoptera: Nitidulidae).

Starvation resistance, or the ability to survive periods without food, can shed light on selection pressure imposed by food scarcity, including chances to invade new regions as a result of human transport. Surprisingly, little information is known about starvation resistance for invasive insect species. Given that native and invasive populations differ in starvation resistance, this would suggest different selection scenarios and adaptive shifts fostering invasion success. Here, we show striking differences in starvation resistance of adult small hive beetles Aethina tumida (SHB) between native and invasive populations. In the laboratory, starvation resistance of freshly emerged laboratory-reared and field-collected adult females and males was evaluated in the beetle's native African range and in their invasive North American range. SHB in their native African range survived longer than SHB in their invasive North American range. Across ranges, females survived longer than males. Field-collected SHB survived in Africa longer than freshly emerged ones, but not in the invasive range. This suggests no selection for starvation resistance in the invasive range, possibly due to differences between African and European-derived honey bee hosts facilitating a trade-off scenario between reproduction and starvation resistance. The ability of adult females to survive up to two months without food appears to be one factor contributing to the invasion success of this species. Assuming food availability is usually high in the invasive ranges, and trade-offs between starvation resistance and fecundity/reproduction are common, it seems as if selection for starvation resistance during transport could set up potential trade-offs that enhance reproduction after invasion. It would be interesting to see if this is a possible general pattern for invasive insect species.

Deformed wings of small hive beetle independent of virus infections and mites.

Insect wing deformities can be caused by viruses, mites and other environmental stressors during development. Here we conducted differential diagnostics of deformed wings in small hive beetles, Aethina tumida (SHB). Adult SHB with and without deformed wings from individual and mass reared scenarios were evaluated for the mite Tyrophagus putrescentiae and for deformed wing virus. Viral load and mite number were similar for SHB with deformed wings and unaffected beetles. Because deformed wings were only observed in individually pupating SHB, a humidity challenge most likely explains the observed clinical symptoms. Our observations support the importance of differential diagnostics.

Bees and flowers: How to feed an invasive beetle species.

Invasive species may exploit a wide range of food sources, thereby fostering their success and hampering mitigation, but the actual degree of opportunism is often unknown. The small hive beetle (SHB), Aethina tumida, is a parasite of honeybee colonies endemic to sub-Saharan Africa. SHBs have now spread on all habitable continents and can also infest colonies of other social bees. To date, the possible role of solitary bee nests as alternative hosts is unknown. Similarly, flowers as possible alternative food sources are not well understood. Here, we show that SHBs can complete an entire life cycle in association with nests of solitary bees Megachile rotundata. The data also show that flowers can serve as alternative food sources. These results support the opportunistic nature of this invasive species, thereby generating further obstacles for mitigation efforts in the field. It also suggests that SHB invasions may result in more serious consequences for endemic bee fauna than previously thought. This provides further motivation to slow down the global spread of this pest, and to improve its management in areas, where it is established.

Sex Ratio of Small Hive Beetles: The Role of Pupation and Adult Longevity.

The sex ratio of sexually reproducing animal species tends to be 1:1, which is known as Fisher's principle. However, differential mortality and intraspecific competition during pupation can result in a biased adult sex ratio in insects. The female-biased sex ratio of small hive beetles (SHBs) is known from both laboratory and field studies, but the underlying reasons are not well understood. Here, we used laboratory mass and individual pupation to test if differential mortality between sexes and/or intraspecific interactions can explain this sex ratio. The data show a significant female-biased adult sex ratio in both mass and individual rearing, even when assuming that all dead individuals were males. Our results therefore suggest that neither differential mortality during pupation nor intraspecific interactions are likely to explain the female-biased sex ratio of freshly emerged adult SHBs. We regard it as more likely that either competition during the larval feeding stage or genetic mechanisms are involved. In addition, we compared our data with previously published data on the sex ratio of both freshly emerged and field-collected SHBs to investigate possible gender differences in adult longevity. The data show a significantly greater female bias in the sex ratio upon emergence, compared to field-collected SHBs, suggesting that adult females have a shorter longevity.