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Honeydew honey is produced by bees from excretions of plant-feeding insects, such as aphids and scale insects. Honeydew on conifers, like fir (Abies alba) or spruce (Picea abies), is produced by different species of the genera Cinara and Physokermes. This means that honeydew honey can stem from different botanical as well as zoological origins, but so far it is not possible to clearly distinguish the different types of honeys. In the attempt to identify distinguishing markers, 19 sugars, 25 amino acids and 9 inorganic ions were quantified in three groups of honeydew honey (fir/Cinara, spruce/Cinara and spruce/Physokermes) with 20 honey samples each. It could be demonstrated that the contents of isomaltose, raffinose, erlose, two undefined oligosaccharides, several amino acids, sulfate, and phosphate differed significantly between the three groups of honey. Furthermore, multivariate analyses resulted in a separation of spruce/Physokermes honey from spruce- or fir/Cinara honey due to its higher contents of phosphate, sulfate, erlose and two undefined oligosaccharides. Moreover, the amino acid composition and the isomaltose as well as the raffinose contents proved useful in the distinction between fir/Cinara and spruce/Cinara honey. In sum, the contents of sugars, amino acids, and inorganic ions in German fir and spruce honeys provide useful information about the botanical and zoological origin of honeydew honeys.

Honeydew honey belongs to the main honey sorts produced in European countries. Phloem sap feeding insects of the order Hemiptera excrete honeydew and honey bees process it into honey. In the case of high osmolality in the phloem sap, hemipteran species counteract osmotic pressure by producing oligosaccharides such as melezitose. Melezitose is a poorly digestible food source for bees, which can lead to honeydew flow disease. Moreover, melezitose-rich honey crystallises quickly and leads to economic losses for beekeepers. To understand the effects of diverse environmental factors on melezitose production, we analysed melezitose proportions with high performance anion-exchange chromatography of 620 honeydew samples collected between 2016 and 2019 considering botanical, zoological, geographical, and meteorological factors. We conducted a two-part logit-model considering all possible variables that could affect melezitose production simultaneously. Higher air temperatures and lower relative humidity levels increased melezitose production. Scale insect species and aphid species on Abies alba produced significantly less honeydew containing melezitose than aphid species on Picea abies. Honeydew with melezitose content was collected more often in natural areas with limited water reservoir capacities. All results lead to the conclusion that in the case the host trees have less access to water and an increasing osmolality of the phloem sap, the melezitose production by hemipteran species is indirectly enhanced. Our results may serve as an indicator of melezitose production and contribute to warning systems for beekeepers that help prevent harmful nutrition for bees or crystallised honey by timely removal of bee colonies from regions at risk.

In general, honey bees (Apis mellifera L.) feed on honey produced from collected nectar. In the absence of nectar, during certain times of the year or in monocultural landscapes, honey bees forage on honeydew. Honeydew is excreted by different herbivores of the order Hemiptera that consume phloem sap of plant species. In comparison to nectar, honeydew is composed of a higher variety of sugars and additional sugars with higher molecular weight, like the trisaccharide melezitose that can be a major constituent of honeydew. However, melezitose-containing honey is known to cause malnutrition in overwintering honey bees. Following the hypothesis that melezitose may be the cause for the so called ‘honeydew flow disease’, three independent feeding experiments with caged bees were conducted in consecutive years. Bees fed with melezitose showed increased food uptake, higher gut weights and elevated mortality compared to bees fed a control diet. Moreover, severe disease symptoms, such as swollen abdomen, abdomen tipping and impaired movement were observed in melezitose-fed bees. 16S-amplicon sequencing indicated that the melezitose diet changed the species composition of the lactic acid bacteria community within the gut microbiota. Based on these results, we conclude that melezitose cannot be easily digested by the host and may accumulate in the hindgut. Within cages or during winter, when there is no opportunity for excretion, the accumulated melezitose can cause severe intestinal symptoms and death of the bees, probably as result of poor melezitose metabolism capabilities in the intestinal microbiota. These findings confirm the causal relation between the trisaccharide melezitose and the honeydew flow disease and indicate a possible mechanism of pathogenesis.