Ecology

Nests
A summary of the nest results can be found in Tables 1 (physical description of nests) and 2 (ploidy). All adult females that were weighed were classed as workers or founder queens using the information found in Rome et al.8, which defined a limit of 593 mg wet weight or 250 mg dry weight to discriminate between workers and founder queens. The average wet weight of founder queens in September was 624 mg (N = 5). All nests recovered had fewer adults present than expected, this was presumed to be due to the loss of adult hornets during the destruction process and subsequent removal of the nest from its original location.
Table 1 Summary of observations from all nests discovered in the UK.
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Table 2 Ploidy of nests.
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Tetbury
The findings from the examination of the Tetbury nest have been described previously10 but are briefly summarised again here. This nest was discovered on 28th September 2016. In total, 70 adult hornets were found in the nest. The wet weight of 57 adult female hornets ranged from 202 to 322 mg with a mean of 256 mg (N = 19), whilst that of 13 adult male hornets ranged from 248 to 326 mg with a mean of 290 mg (N = 7). The nest diameter was 23 cm and the nest contained five combs, four of which contained all life stages (eggs, larvae, pupae, teneral adults, adults) of the Asian hornet. All life stages examined were diploid. The nest was likely derived from a single queen mated to a single drone.
Woolacombe
A nest was discovered in Woolacombe on 27 September 2017. In total, 166 adult hornets were found in the nest, all female. The wet weight of adult female hornets ranged from 172 to 508 mg with a mean of 333 ± 5 mg (N = 166). Based on the information in Rome et al.8, described above, none of the females found in the Woolacombe nest were founder queens. The nest was 27 cm in diameter and is the largest nest discovered in England to date. The nest contained seven combs and all life stages were present, although the larval samples were too degraded for DNA analysis. Haploid individuals were present at the egg and pupal stages, the remainder of the individuals examined were diploid. Based on life stages present and the ploidy, the queen began laying haploid eggs on approximately 2nd September. The genetic analysis (results from COLONY2, verified manually) showed that the offspring sampled were likely to be the product of a single queen and three drones.
Fowey nests 1 + 2
Two nests were discovered in Fowey, Cornwall in 2018 on 3rd and 20th September, 40 m apart. The first nest contained three combs and had a diameter of 15 cm. No eggs or early instar larvae were present but late instar larvae, teneral adults and three adults were present. All individuals sampled were diploid. From the absence of eggs and early instar larvae, it was concluded that the queen was absent/missing in the 2–3 weeks prior to nest discovery. The second nest had a diameter of 19 cm and contained four combs with brood in all stages. Seven adult males and eight females were found in the nest; the males were diploid. All eggs and six of the ten larvae genotyped were haploid, while the pupae, teneral adults and adults were diploid. No queen was found. From the genetic analysis of the two nests, it was shown that both were highly likely to be offspring of a single queen and drone, with the first nest discovered presumably a primary nest and the other nest the secondary nest. From the ploidy of the life-stages present, it was inferred that the queen began laying haploid eggs around the 30th August.
New Alresford nest
The first nest found in Hampshire was discovered in New Alresford on 24th September 2018. The nest was 18 cm diameter and contained four combs with all life stages present. Twenty-eight males and 94 females were found. All the adults, teneral adults and pupae sampled were diploid, while within the larvae, five out of 10 were haploid, and within the eggs, two out of seven were haploid. The queen began laying haploid eggs around the 3rd September. The individuals from this nest were highly likely the offspring of a single queen and two drones.
Brockenhurst nest
The second nest found in Hampshire was discovered in Brockenhurst (approximately 30 miles away from New Alresford) and was destroyed on 04th October 2018. The nest was 18.5 cm diameter and there were three combs present with brood from the larvae stage onwards; no eggs were present indicating a recent loss of the queen or cessation of laying. All larvae and pupae were haploid and adult males were also haploid. The only diploid individuals present were worker females. The queen ceased laying before any diploid (future gyne) eggs were laid. The nest was consistent with being the offspring of a single queen mated with two drones.
Drayton Bassett nest
The first nest of 2019 was discovered on September 2nd at Drayton Bassett, Staffordshire. On arrival at Fera Science Ltd, the nest was too damaged to determine its size. Five adult female hornets were found in the nest. The wet weight of adult female hornets ranged from 197 to 312 mg with a mean of 271.6 (n = 5). The average wet weight of founder queens in September in the study by Rome et al.8 was 624 mg (n = 5). Based on this, it would appear that none of the females found in the nest were founder queens. All life stages were present in the nest, and all individuals genotyped were diploid. The nest was consistent with being the offspring of a single queen mated to a single drone.
Christchurch nests 1 + 2
On 01st October, 2019, a nest 13 cm diameter was discovered in Christchurch, Dorset. Two combs were present in the nest. One adult female hornet was found in the nest. The wet weight of this adult female hornet was 545 mg and the mesoscutum width was 4.6 mm. In a study by Pérez de Heredia et al.16 individuals taken from nests with a unimodal population had one individual per nest that had a mesoscutum width above 4.5 mm; no other individuals in these nests reached a mesoscutum width of 4.5 mm. It is therefore likely that the individual found in this nest was the queen. The combs contained eggs and larvae and had genotypes consistent with being the offspring of the queen that was present. Two eggs were haploid, the remainder of the eggs and larvae genotyped were diploid. On October 10th, a second nest was discovered in Christchurch, 10 m from the first nest, but could not be measured as it was intertwined with vegetation and fragmented upon removal. No adult hornets were found in the nest. Two combs were present, with capped and uncapped cells. Larvae, pupae and teneral adults were found, all of which were diploid. No eggs were found. Both nests from Christchurch were consistent with being the offspring of the same queen, mated to a single drone. The first nest found was likely to be the secondary nest, the second nest found likely to be the primary nest.
Information on all nests is found in Table 1. Map locations for each nest are shown in Fig. 1 and images of each nest are shown in Fig. 2.
Figure 2

Images of UK nests: (a) Tetbury, (b) Woolacombe, (c) Fowey nest 1, (d) Fowey nest 2, (e) Brockenhurst, (f) New Alresford, (g) Drayton Bassett, (h) Christchurch nest 1 and (i) Christchurch nest 2. Where shown, scale bar represents 5 cm.

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Genetic relatedness
Overall, the genetic diversity in the UK is relatively low for all locations, for all three measures used (mean number of alleles per locus, observed and expected heterozygosity; Table 3). However, it should be taken into consideration that the data for each UK nest are from individuals that were all closely related to each other (full, half siblings). A single combined figure for the UK was not calculated as it seems unlikely there is a UK population. Compared to the Asian hornet diversity data from Arca et al. (2015)11, the UK diversity is lower than France, which itself is lower than the diversity found in Asia (Table 2). This trend reflects the likely colonisation history of the hornet, which colonised France from Asia, and the UK incursions are likely to derive from populations on the European mainland.
Table 3 Genetic diversity measure (average number of alleles per locus) and the observed and expected heterozygosity for Asian hornet populations sampled in the British Isles, and from France (data from Arca et al. 201511).
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The occurrence of microsatellite alleles in the UK nests and France and Asia (from Arca et al.11) are given in supplementary material 1. In comparison with the Asian and French data in Arca et al.11, the UK samples had a restricted subset of alleles (35 alleles in total) that were all found in the French populations (60 alleles). In turn, all French alleles were a subset of those found in Asia (178 alleles). Similarly, the majority of private alleles were found in Asia (114), a small number in France (n = 3) and none in the UK (supplementary material 1).
In all cases, individuals recovered near a nest (within 2 km) were offspring of the recovered nearby nest (or nests, where there were primary and secondary nests). The majority of these individuals were found within 500 m of the nest. Most individuals caught in isolated locations away from nests (over 15 km) were not offspring of the recovered nests, with the exception of the individual recovered in Liskeard, which had a genotype compatible with being the offspring of the Fowey nest, some 17 km distant.
To exclude the possibility that any of the founding queens that escaped the destruction of the nest went on to produce viable nests that gave rise to nests caught in the subsequent year, we considered whether the inferred parental genotypes from a nest in year one could be the parents of the inferred parental genotypes in year two. For example, whether the queen from the Tetbury nest in 2016 could have formed a second nest and the offspring from that nest been the parents to the Woolacombe nest in 2017. In no case were the inferred parental genotypes compatible with this scenario. Additionally, where more than one nest was found in a single year (i.e. New Alresford/ Brockenhurst/ Fowey in 2018, Christchurch / Drayton Bassett in 2019), we considered whether the foundress queens and founder drones were full siblings to each other. Again, in no case was this possible (although they could have been half siblings to each other). Genotype data are provided in Supplementary material 2. More

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